A t american museum
JSovitates
PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY
CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024
Number 3512, 40 pp., 11 figures, 1 table May 17, 2006
Redescription of the Cranial Morphology of
Mariliasuchus amarali, and Its Phylogenetic Affinities
(Crocodyliformes, Notosuchia)
HUSSAM ZAHER, 1 DIEGO POL, 2 ALBERTO B. CARVALHO, 1
CLAUDIO RICCOMINI, 3 DIOGENES CAMPOS, 4 AND WILLIAM NAVA 5
ABSTRACT
The cranial morphology of Mariliasuchus amarali, a poorly known notosuchian from the
Late Cretaceous of Southeastern Brazil, is redescribed based on new material. Its phylogenetic
affinities within Crocodylomorpha are evaluated through a parsimony analysis involving 46
taxa and 198 characters. Mariliasuchus is nested well inside the clade Notosuchia, as the sister
group of Comahuesuchus, a derived notosuchian from the Late Cretaceous of Argentina. Both taxa
share the following unambiguous synapomorphies: ventral half of the lacrimal tapering
posteroventrally, not contacting or only slightly contacting the jugal; presence of a large foramen
on the lateral surface of the anterior part of the jugal; presence of procumbent premaxillary and
anterior dentary alveoli; and ectopterygoids that do not participate of the palatine bar. The
presence of procumbent premaxillary teeth, specialized tooth crown morphology, and fore-aft jaw
movements suggests that this group presented complex jaw movements related to specialized
feeding habits.
1 Museu de Zoologia da USP, Servi?o de Vertebrados, Sao Paulo, SP, Brasil (hzaher@ib.usp.br).
2 American Museum of Natural History, Division of Paleontology (dpol@amnh.org).
3 Universidade de Sao Paulo, Instituto de Geociencias, Departamento de Geologia Sedimentar e Ambiental, Sao Paulo,
SP, Brasil (riccomin@usp.br).
4 Departamento Nacional de Produgao Mineral, Museu de Ciencias da Terra, Rio de Janeiro, RJ, Brasil
(dac@abc.org.br).
5 Museu de Paleontologia de Marilia, Marilia, SP, Brasil (willnava@terra.com.br).
Copyright © American Museum of Natural History 2006
ISSN 0003-0082
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AMERICAN MUSEUM NOVITATES
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INTRODUCTION
Sediments of the Bauru Basin, located in
Southeastern Brazil, held one of the most
diverse Upper Cretaceous Crocodyliform fau¬
na known so far (Kellner, 1998). This diversity
is predominantly of Notosuchian taxa, with
some highly specialized forms such as
Sphagesaurus huenei (Price, 1950; Pol, 2003)
and Mariliasuchus amarali (Carvalho and
Bertini, 1999). Despite the exquisite state of
conservation of several fossil crocodilians
unearthed from the Bauru Basin, few have
been described in detail. Although the de¬
scription of Mariliasuchus amarali was
based on a specimen with an almost com¬
plete skull and partially preserved postcra¬
nium, the authors provided little details of
its anatomy. Additionally, the originally de¬
scription is based on a juvenile specimen
(Carvalho and Bertini, 1999). Carvalho
and Bertini (1999) concluded that M. amarali
is more closely related to Notosuchus
terrestris than to any other notosuchian,
allocating it in the family Notosuchidae
along with the genera Notosuchus and
Malawisuchus. We present here a redescription
of the species based on mostly complete skulls
of a subadult and two adult specimens found
in the same locality from which the type
specimen was collected. The new material
described here allows a more careful evalua¬
tion of the phylogenetic affinities of
Mariliasuchus amarali.
GEOLOGICAL SETTING
The fossil remains of Mariliasuchus amarali
reported here as well as the holotype described
by Carvalho and Bertini (1999) were collected
in a road cut at the right margin of the Agua
Formosa creek (coordinates 22°20'28"S and
49°56'46"W), 10 km south from the urban
area of Marilia, about 500 m from the
secondary road between this city and the
locality of Ocaugu (Sao Paulo State), in the
southeastern part of the Bauru Basin. This
basin is a large cratonic depression developed
during the Late Cretaceous in the central-
southeastern portion of the South American
Platform (Fernandes and Coimbra, 1996).
With an area of about 370,000 km 2 (fig. 1A)
and a maximum preserved thickness of 300 m
of clastic deposits, the Bauru Basin contains
a sedimentary sequence—the Bauru Group—
made up mostly of sandy continental deposits,
overlying the Early Cretaceous basaltic lava
flows of the Serra Geral Formation
(Riccomini, 1997). The depocenter of the
basin is coincident with the maximum thick¬
ness of the volcanic pile of the Serra Geral
Formation, in the region of confluence
between the Parana and Paranapanema
rivers (fig. 1A), indicating that the basin
formed by slow and gradual subsidence in
response to the loading of its substrate
(Riccomini, 1997).
The classical stratigraphic division of
the Bauru Group in the state of Sao Paulo
was proposed by Soares et al. (1980) and
encompasses the Caiua (cross-bedded sand¬
stone), Santo Anastacio (massive to slightly
stratified sandstone), Adamantina (massive
to slightly stratified sandstone interlayered
with mudstones), and Marilia (sandstone
and conglomerates with limestone cement)
formations. Suguio (1981) and Fernandes
and Coimbra (2000) also recognized the
Aragatuba (layers of sandstone, siltstone,
and mudstone) and the Presidente Prudente
(sandstone and mudstone) Formations, re¬
spectively.
The stratigraphic relationships of the units
along a WNW-ESE section in the southeast¬
ern part of the basin are depicted in figure IB.
The Caiua and Santo Anastacio formations
are believed to be deposited in a paleodesert,
the former representing the inner part of
a sand sea and the latter the marginal sand
sheets. The Aragatuba Formation probably
represents a paleoswamp (Fernandes et al.,
2003), whereas the Adamantina Formation
comprises sand sheets, loess, and wadi depos¬
its of a peripheral desert (Fernandes and
Coimbra, 2000). The desertic conditions
in this unit decreases upward. Castro et al.
(1999) described lacustrine deposits in the
medium to upper part of the Adamantina
Formation, northeast of the basin depocenter.
Isotopic 5 ls O data from ostracod carapaces
present in these deposits suggest the pre¬
dominance of freshwater conditions (Castro
et al., 1999).
2006
ZAHER ET AL.: CRANIAL MORPHOLOGY OF MARILIASUCHUS AMARALI
3
B WNW
ESE
President© Paidente Fm.
Marftia Fm,
■ Adamantine Fm-, ■
Catua Fm,
Santo Anasiacio Fm.
Arafatuba Fm,
EARLY V v ^ v ^ V Serra Geral Fm.^ V ^ V ^ V ^ V
Fig. 1. A. Location of the Mariliasuchus occurrence in the Bauru Basin (after Riccomini 1997, modified):
1, Precambrian basement rocks; 2, Parana Basin (Ordovician to Triassic); 3, Serra Geral Formation (Early
Cretaceous); 4, Bauru Basin (Late Cretaceous). B. Stratigraphic relationships of the Bauru Group in the
southeastern part of the Bauru Basin: 1, basaltic rocks; 2, cross-bedded sansdstone; 3, massive to slightly
stratified sandstone; 4, massive to slightly stratified sandstone interlayered with mudstones; 5, sandstone,
siltstone, and mudstone; 6, sandstone and mudstone; 7, sandstone and conglomerate with limestone cement;
8, position of Mariliasuchus remains.
4
AMERICAN MUSEUM NOVITATES
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Deposition of the Presidente Prudente
Formation took place in a shallow fluvial
meandering system (Fernandes and Coimbra,
2000) . The Marilia Formation, uppermost
unit of the Bauru Group, represents an influx
of alluvial fans associated with the uplift of the
eastern border of the Bauru Basin (Riccomini,
1997).
The depositional age of the Bauru Group is
still poorly defined. It was first established as
Senonian on the basis of the dinosaur remains
found in both the Adamantina and Marflia
Formations (Huene, 1939). Calcareous micro¬
fossils (ostracods) of the Adamantina
Formation near the city of Presidente
Prudente (western part of the state of Sao
Paulo) were assigned to the Turonian-
Santonian time interval (Dias-Brito et al.,
2001) . The Marilia Formation was considered
of Maastrichtian age, based on vertebrate
fossils (Fernandes and Coimbra, 2000).
Considering the transitional stratigraphic
relationships between the Adamantina and
the overlying Marilia Formation, it is proba¬
ble that the Adamantina Formation is youn¬
ger in the southeastern part of the Bauru
Basin.
The outcrop previously described by
Carvalho and Bertini (1999), and referred to
the Adamantina Formation, is redescribed in
the present work (fig. 2). The exposure
comprises about 1.2 m of brown massive to
slightly stratified mudstone, a loess deposit,
covered by about 3.7 m of massive to slightly
stratified sandstones of sand-sheets. Sand-
sheets include decimetric intercalations of
discontinuous laminae of shale interbedded
with sandstone, which probably corresponds
to ephemeral shallow ponds, and massive or
cross-bedded sandstone with shale intraclasts
representing, respectively, sand-sheet and
ephemeral stream (wadi) with reworked shal¬
low pond deposits. This facies association is
representative of the upper part of the
Adamantina Formation and typical of a pe¬
ripheral desert with sporadic (seasonal?) rain¬
fall and restricted flooding in ephemeral
ponds.
Besides Mariliasuchus, the fossil record in
the sand-sheet facies includes articulated
amphibian remains, eggs, ostracods, ganoid
scales, coprolites, and vertical burrows
(,Skolithos ). Disarticulated vertebrate fossils
(representing remains of Mariliasuchus, un¬
identified fishes, and an undescribed amphib¬
ian) are associated with reworked deposits,
whereas the holotype of Mariliasuchus amarali
was found just above the sedimentary record
of ephemeral ponds.
Based on the stratigraphic position of the
studied section, at the upper part of the
Adamantina Formation (fig. 2), and its re¬
lationship with the overlying Marilia
Formation, it is possible to consider
a Campanian to Maastrichtian age for the
studied samples of Mariliasuchus amarali.
SYSTEMATIC PALEONTOLOGY
CROCODYLIFORMES CLARK, 1986
MESOEUCROCODYLIA WHETSTONE AND
WHYBROW, 1983
Mariliasuchus amarali Carvalho and Bertini,
1999
Figures 3-10
Holotype: Universidade Federal do Rio de
Janeiro, Departamento de Geologia (DG/UFRJ)
50-R, a partially complete articulated individual
with a nearly complete skull and partially
preserved axial and appendicular skeletons.
Referred Specimens: Museu de Zoologia da
Universidade de Sao Paulo (MZSP-PV) 50 and
51, Museu Nacional do Rio de Janeiro (MN)
6298-V and 6756-V (see appendix 1 for in¬
stitutional acronyms used throughout the text).
Locality and Horizon: All the Marilia¬
suchus specimens were collected in a road cut
at the left margin of the Peixe River, 18 km
from the city of Marilia, state of Sao Paulo,
from the upper part of the Adamantina
Formation, Bauru Group, southeastern part
of the Bauru Basin. Based on the stratigraphic
position of the studied section, possibly of
Campanian to Maastrichtian age.
Amended Diagnosis: The lacrimal is barely
exposed on the lateral surface of the snout
(figs. 3, 5), the ectopterygoid contacts the jugal
and the maxilla extensively on its lateral edge,
forming the medial border of at least the last
maxillary alveolus on a ventral plane (fig. 4);
dentary rami forming a mandibular symphysis
that projects anteriorly to form an elongated
2006
ZAHER ET AL.: CRANIAL MORPHOLOGY OF MARILIASUCHUS AMARALI
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AMERICAN MUSEUM NOVITATES
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2006
ZAHER ET AL.: CRANIAL MORPHOLOGY OF MARILIASUCHUS AMARALI
7
and spatulated process with parallel lateral
edges (figs. 6, 7, 8, 9A); there are four pre¬
maxillary, five maxillary, and nine dentary teeth
that are characterized by their heterodonty
(fig. 9); most dentary and maxillary teeth with
anastomosing longitudinal striations formed by
enamel ridges (fig. 10); well-developed serration
of the mesial and distal margins of the crown
(fig. 10A); and one to six longitudinally aligned
tubercles at the base of each crow, which can be
well developed and ornament the whole surface
of the crown base (fig. 10B).
Taxonomic Comments: The name Mari-
liasuchus amaralensis, published in a previous
contribution by Bertini and Carvalho (1999),
is a nomen nudum according to Article 13 of
the International Code of Zoological
Nomenclature (ICZN, 1999), and therefore is
not an available name. Several widely
used taxonomic names are followed through¬
out the text, such as Crocodyliformes, Meso-
eucrocodylia, Neosuchia, and Notosuchia.
Our usage of the first three of them follows
that of Clark (Benton and Clark, 1988; Clark,
1994). Notosuchia was originally created by
Gasparini (1971) to cluster Araripesuchus,
Uruguaysuchus, and Notosuchus. Later, sever¬
al authors pointed out that Sebecosuchia (Pol,
2003; Sereno et al., 2003) was related to this
group, or at least to some of its members
(Buckley et al., 2000; Ortega et al., 2000).
Sereno et al. (2001) defined this taxon using
a stem-based definition of phylogenetic tax¬
onomy. Our results and usage of Notosuchia
is consistent with these propositions.
DESCRIPTION
The material available is composed of four
specimens. Specimen MZSP-PV 50 (figs. 3-8)
consists of an almost complete individual with
skull, mandibles, and most of the postcranial
skeleton. The skull and mandibles are articulated
and crushed on the left side, which caused a slight
displacement of elements on the parasagittal axis
of the skull. The upper right temporal bar is
poorly preserved. Specimen MZSP-PV 51 (fig. 9)
consists of a complete skull and mandibles and the
anterior part of the postcranial skeleton, including
the cervical region, the anterior girdle and
forelimbs, and part of the thoracic region. This
skull is less distorted than MZSP-PV 50, being
slightly crushed dorsoventrally. The right post¬
orbital bar and upper temporal regions are
missing. Specimen MN 6298-V consists of an
almost complete individual with skull, mandible,
and most of the postcranial skeleton. Both skull
and mandible are compressed laterally and
slightly crushed on the sagittal axis. Specimen
MN 6756-V is composed of a partial skull with the
naris, the orbital, and palatal regions preserved,
but lacking the braincase and temporal regions.
The postcranial material of specimens
MZSP-PV 50, 51, and MN 6298-V is currently
under preparation and will be treated else¬
where.
All four skulls analyzed here are oreini-
rostral (sensu Busbey, 1994) with anteriorly
facing external nares, characteristic of terres¬
trial forms. The snout is somewhat constricted
laterally at its midpoint (at the level of the
second maxillary tooth), being more conspic¬
uous in MZSP-PV 50 and MN 6298-V. The
skull widens significantly at the level of the
orbital region. The orbits are large, rounded,
and dorsolaterally exposed when the palpebral
is removed. The temporal region is short and
wide, with large, rounded supratemporal
fossae (sensu Witmer, 1997) occupying most
of the flat skull table. The supratemporal
fossae are separated by a wide flat surface of
the parietal in MZSP-PV 51 and MN 6298-V
(5.92 mm) whereas this separation is markedly
narrow in MZSP-PV 50 (3.33 mm). In MN
6756-V, the parietal is not entirely preserved,
precluding a precise measurement. Onto¬
genetic variation is visible on the shape of
the supratemporal fossae, being more rounded
and wider in the larger specimens whereas they
are more elongated and narrower in the
smaller specimen MN 6298-V. The infratem¬
poral fenestrae are large, triangular, and face
latero-dorsally, being almost completely ex¬
posed on dorsal view. The skull table and
Fig. 3. Skull of MZSP-PV 50 in dorsal view. Scale bar = 1 cm. Anatomical abbreviations are listed
in appendix 1.
AMERICAN MUSEUM NOVITATES
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9
Fig. 5. Skull of MZSP-PV 50 in left lateral view. Scale bar = 1 cm. Anatomical abbreviations are listed
in appendix 1.
Fig. 4. Skull of MZSP-PV 50 in ventral view. Scale bar = 1 cm. Anatomical abbreviations are listed
in appendix 1.
10
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TABLE 1
Measurements of Skulls of MZSP-PV 50, MZSP-
PY 51, MN 6298-V, and MN 6756-V (in millimeters)
MZSP-PV
50
MZSP-PV
51
MN 6298-V MN 6756-V
SAPL
102.19
88.83
72.38
—
MWR
27.30
28.95
16.03
24.11
MLW
15.14
9.95
14.77
MPW
16.80
10.78
—
STW
63.72
41.05
—
SH
—
16.53
17.30
SL
43.72
24.20
31.14
TH
24.76
21.96
—
OL
29.19
21.87
28.05
MLSLo
24.79
18.55
—
MLSLe
13.12
10.47
—
Abbreviations: SAPL skull anteroposterior length;
MWR minimum width of rostrum; MFW minimum
frontal width; MPW minimum parietal width (measured
within the fossae); STW skull table width; SH snout
height at the level of the maximum constriction; SL snout
length from the tip of the snout to the anterior border of
the orbit; TH temporal height measured at the level of the
temporal fossae on their dorsoventral axis; OL orbital
length on its anteroposterior axis; MLSFo maximum
length of supratemporal fossae, measured on its ante¬
roposterior axis; MLSFe maximum length of supratem¬
poral fenestrae, measured on its anteroposterior axis.
dorsal region of the snout of MZSP-PV 51 are
ornamented by grooves and ridges that are
more conspicuous in the region of the snout
than in the temporal region. The other three
specimens do not show the same amount of
ornamentation, being barely ornamented in
most of the dorsal surface of the skull. Cranial
measurements are given in table 1.
Skull Elements
Rostral Region
The premaxillae (figs. 3, 4, 5, 10) form the
ventral and lateral edges of the external nares
and contribute to some degree to the dorsal
edge. The ventral surface of the external nares
bares a small dorsally directed crest formed at
the premaxillary medial suture. This crest
projects anteriorly to form a poorly developed
narrow and pointed anterior process similar to
the condition found in Notosuchus (MACN-
RN 1040). Lateral to the external nares, the
anterior surface of the premaxillae bear
a smooth perinarial depression extending
dorsoventrally. This depression runs parallel
to the narial border, from the level of its
dorsal margin toward the buccal edge. A
similar condition is present in several noto-
suchians (e.g., Comahuesuchus, Simosuchus,
Notosuchus, Baurusuchus ). The lateral surface
of the premaxillae is slightly convex due to the
lodging of the enlarged caniniform root. This
region is deflected and more ornamented
dorsally to the caniniform root whereas the
convex surface tends to be smooth and
vertically oriented. There is at least one large
neurovascular foramen located just anteriorly
to the premaxilla-maxilla suture and another
one at the level of the caniniform tooth.
Contrary to Carvalho and Bertini’s (1999)
interpretation, the premaxilla-maxilla suture
runs posterodorsally to meet the nasal, form¬
ing a blunt posterodorsal process of the
premaxilla that contrasts with the narrow
and pointed condition of Comahuesuchus and
Notosuchus. At the level of the last premaxil¬
lary tooth, the maxilla extends laterally to the
premaxilla, overlapping slightly the latter with
a short anteriorly directed lamina as in
Notosuchus (the condition is unknown in
Comahuesuchus). From that point, the pre¬
maxilla-maxilla suture runs to the palatal
surface, bordering the last alveolus and then
curving gently anteromedially to meet its
counterpart sagitally. In all specimens except
MZSP-PV 50, there is evidence of a median
foramen incisivum between the premaxilla-
maxilla suture at the level of the sagittal plane.
However, it is not possible to clarify the
precise shape and number of foramina due to
the poorly preserved condition of this region
in all specimens. Specimen MZSP-PV 50
seems to lack any trace of foramen incisivum.
The palatal branches of the premaxillae bear
Fig. 6. Mandible of MZSP-PV 50 in dorsal view. Scale bar = 1 cm. Anatomical abbreviations are listed
in appendix 1.
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Fig. 8. Mandible of MZSP-PV 50 in left lateral view. Scale bar = 1 cm. Anatomical abbreviations are
listed in appendix 1.
from 6 to 10 large alveolar foramina on the
anterior part. Each premaxilla bears four
teeth, with the third being caniniform (see
dentition section below).
The maxillae (figs. 3,4, 5, 10) show a tabular
shape and are almost as long as high and
restricted to the dorsolateral and lateral
surfaces of the rostrum. As with the premax¬
illae, the maxillae are vertical and smooth on
their ventral half and deflected with ornamen¬
tations on the dorsal half. Depending on the
specimen, there are 4 to 10 large neurovascular
foramina on the ventral half of the lateral
surface. The suture with the nasal is ante-
roposteriorly directed and runs along almost
all the dorsolateral side of the snout.
Fig. 7. Mandibule of MZSP-PY 50 in ventral view. Scale bar = 1 cm. Anatomical abbreviations are listed
in appendix 1.
14
AMERICAN MUSEUM NOVITATES
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Posteriorly, the maxilla contacts the lacrimal
and participates on the orbital margin, form¬
ing its anteroventral corner. There is no
vestige of an antorbital fenestra. The lateral
contact between the maxilla and the jugal
occurs just below the level of the anterior
margin of the orbit, with the latter sending
a short rounded anterior process laterally to
the maxilla on its dorsal half. The maxilla-
jugal contact runs ventrally toward the buccal
margin where it meets the ectopterygoid. The
buccal margin is straight, lacking the festoon¬
ing pattern present in derived neosuchians.
The palatal branches of the maxillae meet
along their entire length in the sagittal line.
The posterolateral border contacts the ectop¬
terygoid along the medial margin of the last
maxillary tooth (in MZSP-PV 50 and MN
6298-V) or the last two maxillary teeth
(MZSP-PV 51 and MN 6756-V). Medially to
this point, the border of the maxillae runs
transversally and barely enters the anterior
edge of the suborbital fenestrae. The maxilla-
palatine suture extends anteromedially from
the level of the suborbital fenestrae to the
maxillo-palatine foramina where the maxilla
contributes to its anterior edge. The maxilla-
palatine sutures seem to run transversely to
meet at the sagittal plane from the poster¬
omedial edge of the maxillo-palatine forami¬
na. There are five maxillary teeth on each
maxilla that are located in individual alveoli
that are only partially septate (i.e., the in¬
terdental plate is poorly developed). There are
6 to 10 neurovascular foramina along the
lingual edge of the alveoli. In all specimens
both labial and lingual margins are at the
same plane, except in MZSP-PV 51 where the
labial alveolar margin is much more developed
ventrally than the lingual margin.
The pair of nasals (figs. 3, 5) forms most of
the posterodorsal border of the external nares.
They slightly widen posteriorly along their
contact with the premaxillae and maxillae
until reaching the anterior tip of the lacrimal.
The nasal then narrows posteriorly along the
contact with the anterodorsal surfaces of the
lacrimal and prefrontal. The nasal-frontal
suture, positioned just behind the anterior
margin of the orbit, is transversally oriented
and shows clear interdigitation in MZSP-PV
50, MN 6756-V, and MN 6298-V. In MZSP-
PV 51, the precise position of the nasal-
frontal suture cannot be determined due to the
heavy ornamentation.
The lacrimal (figs. 3, 5) is barely exposed on
the lateral surface of the snout and is restricted
to the anterior border of the orbit. It shows
a triangular shape being dorsoanteriorly
expanded at the triple contact between lacri¬
mal, maxilla, and nasal. Ventrally, the lacri¬
mal tapers markedly along the anteroventral
border of the orbit, barely contacting the
expanded anterodorsal border of the jugal
or not contacting at all, as in MN 6298-V.
At that level the lacrimal is embraced medially
by a dorsal flange of the maxilla. The lacrimal
shows a posterior tuberosity exposed dorsally
on its posterodorsal surface that receives the
anterior tip of the palpebral. Dorsally to
this tuberosity, the lacrimal contacts the pre¬
frontal along an anteroposteriorly oriented
suture. This suture continues ventromedially
on the inner surface of the orbit, bordering
the posterior opening of the lacrimal duct,
which is completely included within the
lacrimal.
Both anterior and posterior palpebrals were
preserved in position in specimen MN 6298-V.
Specimens MZSP-PV 50 and 51 retained right
anterior palpebrals in position that were
removed during preparation. A right posterior
palpebral is also preserved in MZSP-PV 50,
but dislocated and laying ventral to the
supratemporal fenestrae along the anterodor¬
sal process of the quadrate (fig. 4). In all
specimens, the anterior palpebrals are blade¬
like, somewhat triangular, elongated, and
strongly curved bones that are latero-poster-
iorly and dorsally positioned to the orbit. The
dorsal surface of the anterior palpebrals is
slightly ornamented. The posterior tip of the
anterior palpebral lies close to the anterior tip
of the posterior palpebral but does not contact
the latter. The posterior palpebrals are short,
broad, and somewhat triangular bones that
lie along the anterior margin of the post¬
orbital.
The prefrontal (figs. 3, 5) is positioned
mostly on the anterodorsal border of the
orbit, running from the dorsal midpoint of the
orbit to the anterodorsal corner, where it
meets the lacrimal. Anteriorly, the prefrontal
expands to meet the nasal on an oblique
2006
ZAHER ET AL.: CRANIAL MORPHOLOGY OF MARILIASUCHUS AMARALI
15
suture. Most of the dorsal surface is orna¬
mented by large rugosities where the palpebral
articulates. The anterodorsal region of the
prefrontal is much wider than the posterior
region, where the prefrontal forms the orbital
margin. Posteromedially, on the inner margin
of the orbit, the prefrontal bears a small
rounded process that overlaps the frontal. The
prefrontal has well-developed prefrontal pil¬
lars strongly sutured to the palate. These are
thin laminae, entirely oriented in an oblique
position. Each prefrontal pillar bears a medi¬
ally oriented process that meets it counterpart
on the sagittal plane, forming a closed bridge
below the olfactory bulbs.
Dorsal and Temporal Region
The frontals (figs. 3, 5) are completely fused
and contribute to most (two-thirds) of the
dorsal margin of the orbit. In all specimens
except MZSP-PV 51, the dorsal surface of the
frontals is flat and barely ornamented. In the
latter, this surface is rugose and bears a slightly
marked longitudinal ridge. A poorly defined
longitudinal ridge is also present in
Comahuesuchus and Notosuchus. It expands
slightly posterolaterally to contact the post¬
orbital in a strait longitudinal suture. The
frontal meets the postorbital and the parietal
at the anterior edge of the supratemporal fossa
in MZSP-PV 50 and MN 6298-V, whereas in
MZSP-PV 51 it seems to expand into the
supratemporal fossa (not visible in MN 6756-
V). The posterior margin of the frontal is
strongly sutured to the parietal via a slightly
concave, transverse, interdigitated suture. The
lateral ventral flange of the frontals forms
a concave and moderately developed dor-
somedial inner orbital wall. Posteriorly, the
flange is sutured to the postorbital and the
dorsal projection of the laterosphenoid.
Anteriorly, the flange is overlapped by a pos¬
terior process of the prefrontal.
The unpaired parietal (fig. 3) forms a dorsal
table that is constricted between the supra¬
temporal fossae, the constriction being wider
in MZSP-PV 51 and MN 6298-V than in the
other two specimens. In MZSP-PV 50, the
parietal table represents only a thin string of
slightly ornamented bone running between the
fossae, whereas in MZSP-PV 51 and MN
6298-V this surface is much broader and
ornamented, being almost as wide as the
frontal at its narrower orbital constriction.
Anteriorly, the parietal contacts the frontal on
a broad and slightly concave suture. The
parietal contacts the postorbital within the
supratemporal fossa. Posterior to this contact,
the parietal forms the entire medial surface of
the supratemporal fossa, being bordered
ventrally by the laterosphenoid and the
ascending anterodorsal process of the quad¬
rate. The laterosphenoid and the quadrate
ascending processes form the medial wall of
the supratemporal fenestra. Lateroposteriorly,
within the supratemporal fossa, the parietal
contacts the squamosal on an interdigitated
suture that passes medially to the anterior
temporal orbital foramen, which is totally
enclosed within the squamosal. Posterior to
this point the parietal meets the supraoccipital
on the dorsal surface of the skull table.
The squamosal (figs. 3, 5) is triradiate, with
the anterior branch contacting the postorbital,
the medial branch contacting the parietal, and
the posterolateral branch contacting the exoc-
cipital and quadrate. In MZSP-PV 50 and
MN 6298-V, the anterior branch is wide
laterally and has a smooth dorsal surface
whereas the medial branch is very narrow and
ornamented dorsally. In MZSP-PV 51, both
branches are ornamented. The postorbital-
squamosal contact is directed posteromedially
from the lateral margin of the skull table to
the lateral margin of the supratemporal fossa,
meeting the anterodorsal process of the
quadrate on a longitudinal suture positioned
at the inner edge of the supratemporal
fenestra. Then, the squamosal suture runs
posteriorly along the inner posterolateral
surface of the supratemporal fossa to meet
the parietal in a triple contact with the latter
and the anterodorsal process of the quadrate.
The lateral margin of the postorbital and
squamosal overhangs the quadrate and quad-
ratojugal, forming a deep otic recess. The
squamosal is extensively sutured to the quad¬
rate within the otic recess, forming the dorsal
and posterior margins of the otic aperture.
The posterolateral branch is almost smooth on
both specimens, showing a median crest that
divides the branch in two distinct lateral
surfaces: one dorsolaterally oriented and the
16
AMERICAN MUSEUM NOVITATES
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other dorsoposteriorly oriented. The dorso-
posterior surface meets the paroccipital pro¬
cess of the exoccipital on a broad contact that
runs mediolaterally along the occipital surface
of the skull.
The postorbital (figs. 3, 4, 5) is also
triradiate, with a narrow medial branch
contacting the frontal and parietal and sepa¬
rating the supratemporal fossa from the orbit,
a posterior branch that contacts the squamo¬
sal, and a descending branch that forms the
dorsal half of the postorbital bar separating
the infratemporal fenestra from the orbit.
Both medial and posterior branches constitute
the anterolateral border of the supratemporal
fossa that is somewhat L-shaped. The ante¬
rolateral border of the postorbital shows
a triangular peglike process forming a step
positioned just below the level of the skull
roof. This process is also present in
Notosuchus, Simosuchus, and Araripesuchus,
where the posterior palpebral articulates. The
posterior branch articulates with the squamo¬
sal dorsally (see description of the squamosal)
and ventrally, within the otic recess. Also
within the otic recess and anteriorly to the otic
notch, the postorbital articulates with the
quadrate and sends a small posteroventral
process that overlaps the ascending process of
the quadratojugal. The ventral branch form¬
ing the dorsal half of the postorbital bar is
smooth and somewhat cylindrical. It embraces
the ascending process of the jugal anteriorly
and contacts in a straight transversally orient¬
ed suture the same process of the jugal
posteriorly.
The jugal (figs. 3, 4, 5) is triradiate, sending
a cylindrical ascending branch directed poster-
odorsally and that meets the postorbital (see
description of postorbital), a broad anterior
branch that contacts the maxilla and ectopter-
ygoid, and a posterior branch that contacts
the quadratojugal. The lateral surface of the
jugal is densely ornamented (except for
the ascending branch) in all specimens. The
anterior branch reaches the level of the
anterior margin of the orbit, where it is
dorsoventrally tall. The anterior edge of the
jugal contacts the maxilla in an extensive and
sinuous suture, overlapping slightly the latter.
The lateral surface of the anterior branch
presents a large foramen that is also present in
Comahuesuchus (as in MOZ P 6131). The
ventral surface of the anterior branch of the
jugal becomes sutured to the extensive pos¬
terolateral process of the ectopterygoid.
Posteriorly, the jugal tapers gradually to form
a thin posterior branch that overlaps laterally
the anterior branch of the quadratojugal. Both
branches contribute to form a dorsoventrally
flattened infratemporal bar.
The quadratojugal (figs. 3, 4, 5) has an
anterior process that forms the posterior half
of the infratemporal bar, lying medial to the
jugal in an extensive longitudinally directed
suture. It expands posteriorly to form a broad
contact with the quadrate and sends a narrow
anterodorsally ascending process that forms
most of the posterior border of the infra¬
temporal fenestra. The ascending process
meets dorsally with the postorbital in a narrow
suture. The posterior and posterodorsal bor¬
ders of the quadratojugal are solidly sutured
to the anterior border of the quadrate. The
posteroventral tip of the quadratojugal does
not reach the articular condyle of the quad¬
rate, thus not contributing to the cranio-
mandibular articulation.
The quadrate (figs. 3, 4, 5) shares most
synapomorphies of the notosuchian clade. The
anterodorsal branch is laterally exposed on
the otic recess whereas the distal body of
the quadrate is projected ventrally at 90° in
respect to the longitudinal axis of the skull.
The distal body of the quadrate is anteropos-
teriorly thin and lateromedially wide and
bears a well-developed ridge on its posterior
surface running from the medial condyle to
the distal tip of the posterolateral process of
the squamosal. A large foramen aereum is
located just medially to this ridge. The distal
body of the quadrate is projected ventrally in
posterior view rather than ventrolaterally as in
most neosuchians. The anterior surface of the
distal body of the quadrate is concave and
smooth, lacking the ridges for the origin of the
adductor bundles. The anterodorsal branch of
the quadrate is broadly exposed laterally on
the otic recess and forms the ventral and
anterior margin of the otic notch. Four
accessory pneumatic foramina are present:
Three are located posteroventrally and one
anteriorly to the otic notch. In addition, there
is a siphoneal foramen located near the
2006
ZAHER ET AL.: CRANIAL MORPHOLOGY OF MARILIASUCHUS AMARALI
17
postorbital-quadrate suture. Anteroventrally,
the pterygoid process of the quadrate is
strongly sutured to the exoccipital, running
anteromedially to meet the basioccipital, basi-
sphenoid, and pterygoid on a longitudinal
somewhat interdigitated suture.
The single supraoccipital (fig. 3) is briefly
exposed on the skull table, representing only
its posterior margin. This exposed part is
lightly ornamented. The supraoccipital is
exposed on the occiput as a triangular surface,
wedging ventrally between the exoccipitals.
The ventral border is excluded from the dorsal
edge of the foramen magnum by the exocci¬
pitals. The occipital surface of the supraocci¬
pital is smooth and bears a sagittal ridge. The
postemporal fenestrae are extremely small and
bordered medially by the supraoccipital,
dorsally by the occipital flange of the squa¬
mosal, and laterally by the exoccipital.
Braincase
The exoccipitals (figs. 3, 4) show two
distinct surfaces: one vertically exposed and
located dorsal to the foramen magnum and
the other one exposed posteroventrally and at
the level of the foramen magnum. Both
surfaces are separated by a transversely ori¬
ented ridge. The vertically oriented surface
extends laterally as the paroccipital process,
which is dorsally and laterally bordered by the
occipital flange of the squamosal and ventrally
by the quadrate. MN 6298-V is a young
individual that shows ontogenetically variable
traits on the occiput. In the latter, the dorsal
surface of the exoccipitals is less concave and
more anteriorly deflected instead of being
vertically oriented as in the larger specimens.
A large cranio-quadrate passage is present
ventral to the paroccipital process and medial
to the triple contact between the squamosal,
quadrate, and paroccipital process. The pos-
teroventral surface extends laterally to meet
the distal body of the quadrate, ventrally to
the cranio-quadrate passage. On its medial
surface, it forms the dorsal and lateral edges of
the foramen magnum and contributes to the
occipital condyle by sending a lateral flange.
Lateral to the foramen magnum and on the
proximal part of the posteroventral surface,
there are five foramina for the exit of the
posterior cranial nerves: The two medial-most
foramina are interpreted to be paired exits for
the Xllth cranial nerve (only visible in MZSP-
PV 50 and 51). Lateral to these there is a larger
foramen subdivided internally by a transverse
wall and a separate smaller, lateral foramen,
which are interpreted as representing the
common passage for the IX, X, and XI cranial
nerves. We failed to find a convincing in¬
terpretation for accommodating the internal
carotid, its course remaining uncertain on
several notosuchian crocodyliforms.
The basioccipital (fig. 4) is exposed poster¬
oventrally in the same plane as the poster¬
oventral surface of the exoccipitals. The
basioccipital forms most of the occipital
condyle and the ventral margin of the foramen
magnum. Anterior to the condyle, the basioc¬
cipital expands laterally, forming a rhomboid
element that laterally contacts the quadrate
and anteriorly the basisphenoid. There is a Y-
shaped, low sagittal crest that diverges ante¬
riorly to surround laterally the foramen
intertympanicum located at the basioccipital-
basisphenoid suture. This sagittal crest is
barely marked in MN 6298-V due to its early
ontogenetic stage. Lateral to this foramen, the
basioccipital-basisphenoid suture is inter¬
rupted by the lateral Eustachian foramen that
is enclosed by these two bones in MZSP-PV
51, whereas in MZSP-PV 50 and MN 6298-V
the Eustachian foramina are enclosed by the
quadrate and basioccipital. In MZSP-PV 50,
the basisphenoid may also have a marginal
participation on the border of the Eustachian
foramina, but this cannot be confirmed un¬
ambiguously on the specimen.
The basisphenoid (fig. 4) is crescentic and
widely exposed on the ventral surface of the
braincase. The lateral margins of the basi¬
sphenoid are bordered by the pterygoid pro¬
cess of the quadrate posteriorly and by the
ascending process of the pterygoid anterolat-
erally and anteriorly. Its ventral surface is flat
near the lateral edges and becomes deeply
concave transversely on its central region. The
concave surface is wide posteriorly and
narrows markedly anteriorly to contact on
a narrow suture the posteromedial surface of
the ascending processes of the pterygoids. The
basisphenoid contacts posteriorly the basioc¬
cipital in a concave suture interrupted by the
18
AMERICAN MUSEUM NOVITATES
NO. 3512
lateral Eustachian foramina, which are located
at the posterolateral end of the bone. The
basisphenoid of Mariliasuchus resembles the
condition present in Sphagesaurus, in which
the bone is widely exposed ventrally with
a slightly concave posterior margin and with
the Eustachian and intertympanicum forami¬
na aligned transversely. This contrasts with
the condition found in Notosuchus, in which
the posterior border is markedly concave and
the foramina are not aligned in the same
transversal plane.
Palate
The palatines (figs. 4, 5) are flat and wide
anteriorly, where they are sutured to each
other, forming the secondary palate, and
narrow and divergent posteriorly, where they
send a barlike process that contacts the
ectopterygoid and pterygoid. The anterior
border of the palatines is positioned slightly
anteriorly to the suborbital fenestrae, suturing
with the maxilla medially to the latter. The
suture is anteromedially directed, reaching the
anterolateral border of the maxillo-palatine
fenestra. Medially to the maxillo-palatine
fenestra, the suture between the palatine and
the maxilla cannot be located with certainty.
The lateral border of the palatine forms the
entire median margin of the elongated sub-
orbital fenestra. At the level of the poster¬
omedial border, the palatines are slightly
deflected ventrally, each one sending a dorso-
ventrally flattened bar in a posterolateral di¬
rection. The posterior half of this bar meets
the medial branch of the ectopterygoid and
the pterygoid flange. The lateral edge of the
posterior half of this process is strongly
sutured to the medial branch of the ectopter¬
ygoid. These two elements overlap and are
sutured to the ventral surface of the pterygoid
flange on its anterolateral region. These
palatine bars form the lateral and anterior
margins of the choanal opening, separating
it from the suborbital fenestra. The palatine
bars are also present in Comahuesuchus,
Notosuchus, and Baurusuchus. In Coma¬
huesuchus and Mariliaschus the palatine bars
stand posteriorly to the suborbital fenestrae,
whereas in Notosuchus and Baurusuchus these
bars are much shorter contacting an ante¬
romedially directed process of the ectopter¬
ygoid at the level of the sub orbital fenestrae,
that is, the latter process of the ectopterygoid
does contribute to the palatine bar in
Notosuchus and Baurusuchus, a very distinct
condition from the one present in Coma¬
huesuchus and Marialiaschus.
As previously described, the ectopterygoid
(Figs. 3, 4) contacts the jugal and the maxilla
extensively on its lateral edge, and on a ventral
plane, forming the medial border of at least
the last maxillary alveolus (two alveoli in
MZSP-PV 51). The contact with the jugal
extends slightly posterolaterally and fails to
project into the medial surface of the post¬
orbital bar, being restricted to the suborbital
region. The lateral edge of the ectopterygoid
contacts the maxilla on its dorsomedial
surface, forming a broad and flat flange of
bone that overlaps the mediodorsal surface of
the maxilla laterally to the sub orbital fenestra.
The medial edge of the ectopterygoid forms
the lateral margin of the sub orbital fenestra,
being slightly constricted at this point. The
posterior end of the ectopterygoid that over¬
laps the ventral surface of the lateral region of
the pterygoid flange is broader and massive
with a striated (rugose) ventral surface. It is
bordered medially by the palatine bar.
The pterygoids (figs. 4, 5) are fused and have
a posterior ascending process that contacts the
basisphenoid and the quadrate on the ventral
and lateral surfaces of the braincase. Medially
and ventrally, the pterygoids narrow markedly,
forming a posterior notch that is continuous to
the medial depression on the ventral surface of
the basisphenoid. The pterygoid flanges are thin
and ventrolaterally oriented, being medially
narrow and expanding laterally to form a large
bladelike process that receives the ectopterygoid
and palatine on the ventral surface of its lateral
edge. Anteriorly, each pterygoid forms a trough¬
shaped choanal groove that projects toward the
palatines. The choanal groove is partially
septated by an anterior process of the pterygoid
that broadens anteriorly to become sutured to
the dorsal surface of the secondary palate.
Dentition
There are four premaxillary, five maxillary,
and nine dentary teeth, which are character-
2006
ZAHER ET AL.: CRANIAL MORPHOLOGY OF MARILIASUCHUS AMARALI
19
ized by their heterodonty (figs. 4, 6, 9).
Specimens MN 6298-V and 6756-V lack most
of their premaxillary teeth, which are well
preserved in MZSP-PV 50 and 51. Maxillary
teeth are preserved in all four specimens,
except in MN 6756-V, which lacks two teeth
on the right row, MZSP-PV 51, which lacks
the last tooth on each row, and MN 6298-V,
which lacks the last tooth of the right row. The
mandible of MN 6756-V is not preserved, but
tooth rows are complete on the dentaries of all
three other specimens, except for MZSP-PV
51 and MN 6298-V, which lack one of the
anteriormost dentary teeth. Except for MN
6298-V, which is a young specimen and for
this reason shows less marked serrations and
thinner enamel coat, all other specimens agree
in most details. The specimens MZSP-PV 50
and 51 have preserved better dentition; there¬
fore we will focus our detailed description
below on these two specimens.
All teeth are covered by a thick layer of
enamel that is preserved with a dark-brown
coloration (figs. 9, 10). The two anteriormost
premaxillary teeth are facing anteroventrally
rather than ventrally as in other crocodyli-
forms (fig. 9A). These elements are smooth,
conical, and gently curved downward. The
third premaxillary tooth is caniniform and
conical, being smooth in MZSP-PV 51 but
showing clear longitudinal striation all on
its surface in MZSP-PV 50 (fig. 9B). The
fourth premaxillary tooth is the smallest of
the four, showing a blunt but conical shape
with barely developed longitudinal striations.
The general shape of all five maxillary teeth
is of a blunt aspect, with the labial side being
more convex than the lingual side. The first
maxillary tooth is similar in size to the last
premaxillary tooth. The three following max¬
illary teeth are larger, whereas the last one is
reduced (being the smallest element of the
maxillary tooth row). They show well-de¬
veloped serrations on the mesial and distal
margins of the crown, except for the first
tooth, which lacks a well-developed serration
on its mesial margin. The serration is com¬
posed of a series of rounded tubercles,
instead of the sharp denticles present in
ziphodont crocodyliforms (see Prasad and de
Broin, 2002). These tubercles are mainly
formed by the thickening of the enamel coat.
All the maxillary teeth have anastomosing
longitudinal striations formed by enamel
ridges. At the base of each crown, some of
these ridges bear one to six longitudinally
aligned tubercles similar to the ones that form
the marginal serration. These tubercles can be
well developed and ornament the whole
surface of the crown base, being more
conspicuous at the third and fourth maxillary
teeth (fig. 10B). The distal serrated margin
of the second and third maxillary teeth is
directed posteromedially rather than poster¬
iorly (a condition more developed in MZSP-
PV 50). Both specimens show extensive wear
facets on the lingual surface of some of their
maxillary teeth. In MZSP-PV 50, the wear
facet is located apically on the lingual surface,
being extended both anteriorly and posteriorly
in the second right and fourth teeth on both
sides. The second left maxillary tooth shows
a wear facet that extends more posteriorly and
basally than in the other teeth. This latter
pattern is also present in the second right,
third, and fourth teeth of MZSP-PV 51. The
second left tooth of MZSP-PV 51 bears
a reduced wear facet that is located antero-
apically. Where visible in the wear facet, the
striae are directed obliquely from the ante-
rodistal margin of the wear facet toward its
posterobasal margin. Such striae suggest
a significant anteroposterior movement of
the lower jaws, which is congruent with the
presence of an anteroposteriorly enlarged
articular surface for the quadrate condyles
(see description of the mandible). The two
anteriormost teeth of the dentary are conical
and directed horizontally in a markedly pro¬
cumbent position. Posteriorly along the tooth
row, the mandibular teeth shift progressively
to a more erect position. The third and fourth
dentary teeth are conical and have slightly
serrated distal and mesial margins, being still
directed anterodorsally. Posterior to this
point, the remaining dentary teeth are erect
and blunt shaped. The size and shape of these
crowns as well as the enamel surface closely
match the morphology of their corresponding
maxillary teeth. The wear facets are well
developed and visible only on the sixth to
eighth dentary teeth. These wear facets are
located on the labial surface of the crowns and
perfectly match the shape and extension of the
20
AMERICAN MUSEUM NOVITATES
NO. 3512
Fig. 9. A. Dorsal view of
mandibular symphysis of MZSP-
PV 51. B. Ventral view of ante¬
rior palatal region of the skull of
MZSP-PV 51.
2006
ZAHER ET AL.: CRANIAL MORPHOLOGY OF MARILIASUCHUS AMARALI
21
22
AMERICAN MUSEUM NOVITATES
NO. 3512
wear facets present in the corresponding
maxillary teeth, implying that a pattern of
tooth-to-tooth occlusion was present in
Mariliasuchus amarali. Such a pattern was
only found previously in Sphagesaurus huenei
among crocodyliforms (Pol, 2003).
To accommodate the highly autapomorphic
position of the two anteriormost teeth, the
corresponding alveoli of both premaxillae and
dentaries are in an almost horizontal position,
with an enlarged labial wall and a highly
reduced lingual wall.
Mandible
The complete mandible is preserved in
MZSP-PV 50 (figs. 6, 7, 8), MZSP-PV 51,
and MN 6298-V. The dentaries are dorsoven-
trally low and laterally convex at the mandib¬
ular symphysis. The dentary rami converge to
meet and form a mandibular symphysis that
tapers anteriorly along its posterior half.
Anteriorly, the symphyseal region is elongated
and spatulated, with parallel lateral edges. At
this point, the symphyseal region has a pecu¬
liar, large, and flattened dorsal surface be¬
tween the two parallel tooth rows. In all three
specimens the ventral surface of the symphysis
is ornamented, whereas the lateral surface is
smooth and bears several large neurovascular
foramina, varying from four to eight. Just
posterior to the mandibular symphysis, the
dentaries diverge and the tooth rows (last four
teeth) are medially inset on their ramus, being
lingually bordered by a thin lamina of the
splenial. The lateral surface of the posterior
half of the dentary is mostly smooth and
lateromedially broad in the larger specimens.
The dorsal posterior process of the dentary is
forked, receiving an acute anterior process of
the surangular in an interdigitated suture. The
ventral branch of the dorsal posterior process
forms the anterodorsal margin of the external
mandibular fenestra, and contacts the angular
at the anterior edge of the mandibular
fenestra. The ventral posterior process of the
dentary is extremely reduced and fails to
extend underneath the mandibular fenestra,
the angular-dentary suture being directed
dorsoventrally.
The splenials (fig. 7) represent thin laminae
that form the posterior third of the dorsal
surface of the mandibular symphysis. The
splenial participation on the ventral surface of
the symphysis is much more restricted, con¬
stituting only the posterior tip of the symphy¬
sis ventrally, where it forms a peglike pro¬
tuberance on the sagittal midline. The poste¬
rior surface of the symphysis is notably high
and is entirely formed by the splenials, which
contact each other on a solid interdigitated
suture. Posterior to the symphysis, the sple¬
nials contribute to the ventral surface of the
mandibular rami on their anterior half,
curving gently dorsally and being restricted
to the medial surface of the rami on their
posterior half. The medial surface of the
mandibular rami is covered by a flat lamina
of the splenial that bears a moderately large
foramen intramandibularis oralis posterior to
the mandibular symphysis, located at the level
of the seventh dentary tooth. The posterior
edge of the splenials forms the anterior border
of the internal mandibular fenestra. Its poster-
oventral margin contacts the anterior process
of the angular that wedges between the
dentary and splenial.
The surangular (figs. 6, 8) is an elongated
bone that forms the dorsal margin of the
posterior half of the mandibular rami. The
anterior process is forked like the posterior
dorsal process of the dentary with which it
interdigitates. The medial surface of the
anterior branch extends anteriorly between
the dorsal margin of the splenial and the
dentary, reaching the lateral border of the last
dentary alveolus. At the level of the ante¬
rodorsal border of the inner mandibular
fenestra, the anterior process of the surangular
has a concave and spatulate rugose surface
that projects medially and ventrally. This
surface might represent the coronoid bone
Fig. 10. A. Left lateral view of the skull of MZSP-PV 50 showing the buccal side of the premaxillary and
maxillary tooth rows. B. Detail of the second maxillary tooth of MZSP-PV 51 in lingual view. Scale bar =
1 cm.
2006
ZAHER ET AL.: CRANIAL MORPHOLOGY OF MARILIASUCHUS AMARALI
23
that fused to the surangular. However, there is
no unambiguous evidence that the coronoid
was present in any of the examined specimens.
Posterior to this spatulate process, the sur¬
angular broadens markedly along its contact
with the lateral surface of the articular. The
surangular projects a ventrally directed point¬
ed process that forms the posterior margin of
the external mandibular fenestra. This process
is mostly overlapped laterally by the angular.
The surangular forms the upper lateral surface
of the postdentary region of the mandible,
joining the angular in a straight suture
oriented anteroposteriorly. The posterior tip
of the surangular reaches the dorsolateral
surface of the retroarticular process, covering
it partially.
The angular (fig. 7) forms the ventral
margin of the postdentary ramus of the
mandibles as well as the ventral half of their
lateral surface, posterior to the external
mandibular fenestra. Its anterior part forms
the entire ventral margin of the mandibular
fenestra and wedges anteriorly between the
splenial and the dentary, forming a pointed,
anteriorly directed process on the ventral
surface of the mandible. The lateral lamina
of the angular also forms most of the anterior
margin of the external mandibular fenestra,
being overlapped by the dentary anteriorly to
this region. Posterior to the mandibular
fenestra, the angular is smooth, extending
caudally to overlap laterally the ventrolateral
surface of the articular and retroarticular
process.
The articular (fig. 6) is roughly triangular,
having an acute tip that extends anteriorly to
the articular facet for the quadrate, deflecting
ventrally, being bordered laterally by the
surangular and fitting into the U-shaped
angular surface located posteriorly and medi¬
ally to the mandibular fenestra. The facet is
anteroposteriorly long and broadens poster¬
iorly. Its dorsal surface is convex, lacking
a well-developed longitudinal ridge that fits
between the quadrate condyles. The ventral
surface of the articular facet projects medially
as a shelf, overhanging the ventral surface of
the angular. The retroarticular process (figs. 6,
7, 8) has a broad, rounded, and slightly
concave surface that projects posteroventrally
from the articular facet.
DISCUSSION
Assignment of Specimens to
MARILIASUCHUS AMARALI
The four specimens described herein share
all the diagnostic features present in the type
specimen and exemplified in the Amended
Diagnosis. However, the former specimens
differ from the latter in numerous aspects that
are ontogenetically related, since the type
specimen is a juvenile whereas the material
described here consists of two subadults and
two adults. Specimens MN 6298-V, MN 6756-
V, and MZSP-PV 50 are very similar in most
aspects, the former representing a younger
adult. Although MZSP-PV 51 shares all
autapomorphic traits of the species, it shows
several significant morphological differences,
such as the presence of a foramen incisivum
(absent in the other three specimens), a dense
ornamentation of the skull table and dorsal
region of the snout (only poorly ornamented
in MN 6298-V, MN 6756-V, and MZSP-PV
50), a wider parietal width between the
supratemporal fossae, and the presence of
a longitudinal ridge on the frontal (absent in
MN 6298-V, MN 6756-V, and MZSP-PV 50).
These differences might suggest that MZSP-
PV 51 belongs to a different species from the
other three specimens. Alternatively, these
differences may be related to sexual dimor¬
phism or individual variation. Here MZSP-PV
51 is referred to Mariliasuchus amarali, pro¬
visionally accepting the latter hypotheses to
explain the observed differences. However,
more material is needed in order to clarify this
issue.
Phylogenetic Affinities of
Mariliasuchus amarali
To test the phylogenetic affinities of
Mariliasuchus amarali , we used an extended
version of the data matrix furnished by Pol
and Norell (2004b), including all notosuchian
taxa relevant for the analysis (see appendices 2
and 3). A total of 46 taxa were scored for 198
characters. Parsimony analysis, using PAUP
4.0 (beta 10) (Swofford, 2003) with a heuristic
search strategy (100 replicates of Wagner trees
followed by TBR branch swapping, resulted in
12 most parsimonious trees with 658 steps (Cl
24
AMERICAN MUSEUM NOVITATES
NO. 3512
CROCODYLIFORMES
/
NOTOSUCHIA
/
NEOSUCHIA
/
- Gracilisuchus
- Terrestrisuchus
- Dibothrosuchus
- Orthosuchus
_ Kayenta form
- Protosuchus
. Hemiprotosuchus
- Zaraasuchus
- Gobiosuchus
-Zosuchus
- Sichuanosuchus
- Shantungosuchus
- Fruita form
- Flsisosuchus
- Araripesuchus gomesi
- Araripesuchus patagonicus
- Uruguaysuchus
- Simosuchus
- Malawisuchus
- Candidodon
- Notosuchus
- Comahuesuchus
- Mariliasuchus
- Chimaerasuchus
- Sphagesaurus
- Baurusuchus
- Bretesuchus
- Iberosuchus
- Lomasuchus
- Peirosaurus
■ Theriosuchus
- Aliigatorium
- Pholidosaurus
- Sokotosuchus
- Dyrosaurus
- Metriorhynchus
- Peiagosaurus
- Steneosaurus
- Goniopholis
- Eutretanonew
- Bernissartia
- Hylaeochampsa
- Boreaiosuchus
- Gaviatis
- Crocodylus
-Alligator
2006
ZAHER ET AL.: CRANIAL MORPHOLOGY OF MARILIASUCHUS AMARALI
25
= 0.36; RI = 0.67), the strict consensus of
which is presented in figure 11. The 12
phylogenetic hypotheses differ in the relation¬
ships of some neosuchian crocodyliforms
(e.g., Hylaeochampsa, Borealosuchus, and
Pholidosaurus) and the alternative position of
Notosuchus either as the sister group of clade 6
or clade 7 within Notosuchia (or Ziposuchia
sensu Ortega et al., 2000). In contrast to some
previous phylogenetic analyses (e.g., Clark,
1994; Buckley et al., 2000; Ortega et al., 2000;
Turner, 2004), the genus Araripesuchus ap¬
pears as the basalmost notosuchian instead of
being a basal member of Neosuchia, as in
a recent analysis (Pol and Apesteguia, 2005).
According to the present analysis,
Mariliasuchus appears nested well inside
the clade Notosuchia, as the sister group
of Comahuesuchus, a derived and highly
autapomorphic notosuchian from the Late
Cretaceous of Argentina. These two taxa
share four unambiguous synapomorphies:
the ventral half of the lacrimal tapering
posteroventrally, not contacting or only slight¬
ly contacting the jugal (character 192 [1]), the
presence of a large foramen on the lateral
surface of the anterior part of the jugal
(character 193 [1]), the presence of procum¬
bent premaxillary and anterior dentary alveoli
[character 194 (1)], and ectopterygoids that do
not participate of the palatine bar (character
196 [0]). Character 194 confers to both
Mariliasuchus and Comahuesuchus an unusual
condition in which the anteriormost (pre¬
maxillary and dentary) teeth are set horizon¬
tally instead of vertically. These peculiar
characteristics, related to specialized dental
crown morphology and the presence in de¬
rived notosuchians of fore-aft jaw move¬
ments, suggests that this group presented
complex jaw movements related to specialized
feeding habits still poorly understood for the
majority of the taxa.
Jacobs et al. (1990) and, more recently,
Nobre and Carvalho (2002) suggested that
Candidodon itapecuruense from the Lower
Cretaceous Parnaiba basin is more closely
related to Malawisuchus than to any other
notosuchian, due to their similarities in dental
morphology (i.e., lingual base of the crown
ornamented with a cuspidate cingulum; Clark
et al., 1989; Gomani, 1997; Carvalho and
Bertini, 2000). Similarly, one could interpret
the complex lingual cuspidate ornamentation
at the base of the crown of Mariliasuchus as
homologous to the lingual cuspidate cingula in
the teeth of Candidodon and Malawisuchus.
However, such similarity may well be superfi¬
cial, as recent works documented a high di¬
versity of complex crown morphologies among
crocodyliforms (Clark et al., 1989; Wu and
Sues, 1996; Gomani, 1997; Wu et al., 1997;
Larsson and Sidor, 1999; Buckley et al., 2000;
Pol, 2003). The present analysis supports the
hypothesis previously proposed by Jacobs et al.
(1990) and Nobre and Carvalho (2002), with
Candidodon and Malawisuchus forming a clade
(4) supported by five ambiguous synapomor¬
phies (9 [1], 122 [0], 140 [0], 149 [0], 161 [2];
contra Carvalho et al., 2004).
Although the recently described notosu¬
chian Anatosuchus minor was hypothesized to
be the sister group of Comahuesuchus (Sereno
et al., 2003), it was not included in this
analysis, awaiting a more detailed description.
However, Sereno et al.’s (2003) prelimi¬
nary description of the skull shows that
Anatosuchus may lack the synapomorphies
supporting clade 6 (fig. 11), suggesting
that it might not represent the sister group
of Comahuesuchus. A more detailed phylo¬
genetic analysis including Anatosuchus, as
well as Comahuesuchus, Mariliasuchus, and
Candidodon, is needed before any more
accurate taxonomic or biogeographic conclu¬
sions concerning these taxa can be drawn.
Fig. 11. Strict consensus of the 12 most parsimonious topologies that resulted from a strict parsimony
analysis using PAUP 4.0 (beta 10). Unambiguous synapomorphies for the labeled nodes are: Node 1: 95 (0),
104 (2), 151 (1). Node 2: 1 (1), 74 (1), 79 (0), 106 (1). Node 3: 78 (1), 107 (1), 141 (1). Node 4: 140 (0). Node 5:
195 (1), 198 (1). Node 6: 192 (1), 193 (1), 194 (1), 196 (0). Node 7: 121 (1), 130 (1), 134 (1), 148 (1). Node 8:
105 (3), 124 (1). Node 9: 3 (0), 9 (2), 79 (1), 80 (1), 106 (0), 118 (1), 120 (0), 128 (0), 155 (0), 158 (1).
26
AMERICAN MUSEUM NOVITATES
NO. 3512
ACKNOWLEDGMENTS
The authors thank M. Norell (American
Museum of Natural History, New York), M.
Wilkinson (The Natural History Museum,
London), F. de Broin, P. Taquet, C. de Muizon
(Museum National d’Histoire Naturelle de
Paris), J. Bonaparte, A. Kramarz (Museo
Argentino de Ciencias Naturales), J. Clark
(George Washington University), S.A.K. de
Azevedo (Museu Nacional do Rio de Janeiro),
and I.S. Carvalho (Universidade Federal do
Rio de Janeiro) for permission to analyze speci¬
mens under their care. Jim Clark and Chris
Brochu provided thoughtful comments that
improved the quality of the manuscript. We also
deeply thank Pablo Goloboff for his help, advice,
and assistance in the computer-assisted phyloge¬
netic analysis using PAUP*. The present contri¬
bution benefited from grants of FAPESP (01/
00162-3) and CNPq (303413/2002-6) to the
senior author.
REFERENCES
Benton, M.J., and J. Clark. 1988. Archosaur
phylogeny and the relationships of the
Crocodylia, In M.J. Benton (editor), The
phylogeny and classification of the tetrapods.
Volume 1: amphibians, reptiles, birds. The
Systematics Association Special Volume 35A:
295-338.
Bertini, R.J., and I.S. Carvalho. 1999. Distribuigao
cronologica dos crocodilomorfos notossuquios
e ocorrencias nas bacias cretacicas brasileiras.
Boletim do 5°. Simposio sobre o Cretaceo do
Brasil / ler Simposio sobre el Cretacico de
America del Sur, Rio Claro, Universidade
Estadual Paulista, 1999): 517-523.
Brochu, C.A. 1997. Fossils, morphology, diver¬
gence timing, and the phylogenetic relation¬
ships of Gavialis. Systematic Biology 46:
479-522.
Buckley, G.A., and C.A. Brochu. 1999. An
enigmatic new crocodile from the Upper
Cretaceous of Madagascar. In D.M. Unwin
(editor), Special Papers in Palaeontology 60:
149-175.
Buckley, G.A., C.A. Brochu, D.W. Krause, and
D. Pol. 2000. A pug-nosed crocodyliform from
the Late Cretaceous of Madagascar. Nature
405: 941-944.
Busbey, A.B. 1994. Structural consequences of skull
flattening in crocodilians. In J. Thomason
(editor), Functional morphology and verte¬
brate paleontology: 173-192. Cambridge:
Cambridge University Press.
Buscalioni, A.D., and J.L. Sanz. 1988. Phylogenetic
relationships of the Atoposauridae (Archo-
sauria, Crocodylomorpha). Historical Biology
1: 233-250.
Carvalho, I.S., and R.J. Bertini. 1999. Mariliasuchus :
um novo Crocodylomorpha (Notosuchia) do
Cretaceo da Bacia Bauru. Geologia Colombiana
24: 83-105.
Carvalho, I.S., and R.J. Bertini. 2000. Contexto
geologico dos notossuquios (Crocodylo¬
morpha) cretacicos do Brasil. Geologia
Colombiana 25: 163-184.
Castro, J.C., D. Dias-Brito, E.A. Musacchio,
J.M. Suarez, M.S.A.S. Maranhao, and
R. Rodrigues. 1999. Arcabougo estratigrafico
do Grupo Bauru no Oeste Paulista. Boletim do
5°. Simposio sobre o Cretaceo do Brasil / ler
Simposio sobre el Cretacico de America del
Sur, Rio Claro, Universidade Estadual Paulista
(1999): 509-515.
Clark, J.M. 1994. Patterns of evolution in Mesozoic
crocodyliformes. In N.C. Fraser and H.-D. Sues
(editors), In the shadow of dinosaurs: 84-97.
Cambridge: Cambridge University Press.
Clark, J.M., L.L. Jacobs, and W.R. Downs. 1989.
Mammal-like dentition in a Mesozoic crocody-
lian. Science 244: 1064-1066.
Dias-Brito, D., E.A. Musachchio, J.C. Castro,
M.S.A.S. Maranhao, J.M. Suarez, and R.
Rodrigues. 2001. Grupo Bauru: uma unidade
continental do Cretaceo no Brasil—concepgoes
baseadas em dados micropaleontologicos,
isotopicos e estratigraficos. Revue de
Paleobiologie 20: 245-304.
Fernandes, L.A., and A.M. Coimbra. 1996. A
Bacia Bauru (Cretaceo Superior, Brasil). Anais
da Academia Brasileira de Ciencias 68:
195-205.
Fernandes, L.A., and A.M. Coimbra. 2000.
Revisao estratigrafica da parte oriental da
Bacia Bauru (Neocretaceo). Revista Brasileira
de Geociencias 30: 717-728.
Fernandes, L.A., P.C.F. Giannini, and A.M. Goes.
2003. Aragatuba Formation: palustrine depos¬
its from the initial sedimentation phase of the
Bauru Basin. Anais da Academia Brasileira de
Ciencias 75: 173-187.
Frey, E. 1988. Das Tragsystem der Krocodile—eine
biomechanische und phylogenetische Analyse.
Stuttgarter Beitrage zur Naturkunde
Serie A (Biologie) 426: 1-60.
Gasparini, Z.B. 1971. Los Notosuchia del Cretacico
de America del Sur como um nuevo infraorden
de los Mesosuchia (Crocodylia). Ameghiniana
8: 83-103.
2006
ZAHER ET AL.: CRANIAL MORPHOLOGY OF MARILIASUCHUS AMARALI
27
Gasparini, Z.B., L.M. Chiappe, and M. Fernandez.
1991. A new Senonian Peirosaurid (Crocody-
lomorpha) from Argentina and a synopsis of
the South American Cretaceous crocodilians.
Journal of Vertebrate Paleontology 11: 316—
333.
Gomani, E. 1997. A crocodyliform from the Early
Cretaceous dinosaur beds, northern Malawi.
Journal of Vertebrate Paleontology 17:
280-294.
Huene, F. 1939. Carta de F. von Huene ao Dr.
Euzebio de Oliveira. Mineragao e Metalurgia 4:
190.
ICZN. 1999. International Code of Zoological
Nomenclature, 4th ed. London: International
Trust of Zoological Nomenclature.
Jacobs, L.L., D.A. Winkler, Z.M. Kaufulu, and
W.R. Downs. 1990. The dinosaur beds of
northern Malawi, Africa. National Geographic
Research 6(2): 162-204.
Kellner, A.W.A. 1998. Panorama e perspectiva do
estudo de repteis fosseis no Brasil. Anais da
Academia Brasileira de Ciencias 70(3):
647-676.
Larsson, H.C.E., and C.A. Sidor. 1999. Unusual
crocodyliform teeth from the Late Cretaceous
(Cenomanian) of southeastern Morocco.
Journal of Vertebrate Paleontology 19:
398^401.
Nobre, P.E., and I.S. Carvalho. 2002. Osteologia
do cranio de Candidodon itapecuruense
(Crocodylomorpha, Mesoeucrocodylia) do
Cretaceo do Brasil. Boletim do 6°. Simposio
sobre o Cretaceo do Brasil / 2nd. Simposio
sobre el Cretacico de America del Sur (2002):
77-82.
Ortega, F., A.D. Buscalioni, and Z.B. Gasparini.
1996. Reinterpretation and new denomination
of Atacisaurus crassiproratus (Middle Eocene;
Issel, France) as cf. Iberosuchus (Croco¬
dylomorpha: Metasuchia). Geobios 29:
353-364.
Ortega, F., Z.B. Gasparini, A.D. Buscalioni, and
J.O. Calvo. 2000. A new species of
Araripesuchus (Crocodylomorpha, Mesoeu¬
crocodylia) from the lower Cretaceous of
Patagonia (Argentina). Journal of Vertebrate
Paleontology 20: 57-76.
Osmolska, H., S. Hua, and E. Buffetaut. 1997.
Gobiosuchus kielanae (Protosuchia) from the
Late Cretaceous of Mongolia: anatomy and
relationships. Acta Paleontologica Polonica 42:
257-289.
Pol, D. 1999a. El esqueleto postcraneano de
Notosuchus terrestris (Archosauria: Croco-
dyliformes) del Cretacico Superior de la
Cuenca Neuquina y su informacion filogeneti-
ca. Tesis de Licenciatura, Facultad de Ciencias
Exactas y Naturales, Universidad de Buenos
Aires, Argentina, 158 pp.
Pol, D. 1999b. Basal mesoeucrocodylian relation¬
ships: new clues to old conflicts. Journal of
Vertebrate Paleontology 19(suppl. to no. 3):
69A.
Pol, D. 2003. New remains of Sphagesaurus huenei
(Crocodylomorpha: Mesoeucrocodylia) from
the Late Cretaceous of Brazil. Journal of
Vertebrate Paleontology 23: 817-831.
Pol, D., and S. Apesteguia. 2005. New
Araripesuchus remains from the Early Late
Cretaceous (Cenomanian-Turonian) of Patagonia.
American Museum Novitates 3490: 1-38.
Pol, D., and M.A. Norell. 2004a. A new crocodyli¬
form from Zos Canyon Mongolia. American
Museum Novitates 3445: 1-36.
Pol, D., and M.A. Norell. 2004b. A new gobio-
suchid crocodyliform taxon from the
Cretaceous of Mongolia. American Museum
Novitates 3458: 1-31.
Prasad, G.V.R., and L. de Broin. 2002. Late
Cretaceous crocodile remains from Naskal
(India): comparisons and biogeographic affin¬
ities. Annales de Paleontologie 88(1): 19-71.
Price, L.I. 1950a. On a new crocodilian,
Sphagesaurus , from the Cretaceous of the
State of Sao Paulo, Brazil. Anais da
Academia Brassileira de Ciencias 22(1): 77-85.
Riccomini, C. 1997. Arcabougo estrutural e aspec-
tos do tectonismo gerador e deformador da
Bacia Bauru no Estado de Sao Paulo. Revista
Brasileira de Geociencias 27: 153-162.
Sereno, P.C., H.C.E. Larsson, C.A. Sidor, and
B. Gado. 2001. The giant crocodyliform
Sarcosuchus from the Cretaceous of Africa.
Science 294: 1516-1519.
Sereno, P.C., C.A. Sidor, H.C.E. Larsson, and
B. Gado. 2003. A new notosuchian from the
Early Cretaceous of Niger. Journal of
Vertebrate Paleontology 23: 477-482.
Soares, P.C., P.M.B. Landim, V.J. Fulfaro, and
A.F. Sobreiro Neto. 1980. Ensaio de caracter-
izagao estratigrafica do Cretaceo no Estado de
Sao Paulo: Grupo Bauru. Revista Brasileira de
Geociencias 10: 177-185.
Suguio, K. 1981. Fatores paleoambientais e paleo-
climaticos e subdivisao estratigrafica do Grupo
Bauru. In A Formagao Bauru no Estado de Sao
Paulo e regioes adjacentes: 15-26. Sao Paulo:
Sociedade Brasileira de Geologia.
Swofford, D.L.PAUP*. Phylogenetic analysis using
parsimony (*and other methods). Version 4.
Sunderland, MA: Sinauer Associates.
Turner, A.H.T. 2004. Crocodyliform biogeography
during the Cretaceous: evidence of Gondwanan
vicariance from biogeographical analysis.
28
AMERICAN MUSEUM NOVITATES
NO. 3512
Proceedings of the Royal Society London B
271: 2003-2009.
Witmer, L.M. 1997. The evolution of the antorbital
cavity of archosaurs: a study in soft-tissue
reconstruction in the fossil record with an
analysis of the function of pneumaticity.
Journal of Vertebrate Paleontology, Memoir
3: 1-73.
Wu, X.-C., and H.-D. Sues. 1996. Anatomy and
phylogenetic relationships of Chimaeresuchus
paradoxus, an unusual crocodyliform reptile
from the Lower Cretaceous of Hubei, China.
Journal of Vertebrate Paleontology 16: 688—
702.
Wu, X.-C., H.-D. Sues, and Z.-M. Dong. 1997.
Sichuanosuchus shuhanensis : a new ?Early
Cretaceous protosuchian (Archosauria:
Crocodyliformes) from Sichuan (China), and
the monophyly of Protosuchia. Journal of
Vertebrate Paleontology 17: 89-103.
APPENDIX 1
APPENDIX 2
Institutional Acronyms and
Anatomical Abbreviations
Character List Corresponding to Data
Matrix Used in Phylogenetic Analysis
Institutional
DG/UFRJ Departamento de Geologia,
Universidade Federal do Rio de
Janeiro, Brazil.
MZSP-PV Museu de Zoologia, Universidade
de Sao Paulo, Brazil.
MN Museu Nacional, Universidade
Federal do Rio de Janeiro, Brazil.
MACN Museo Argentino de Ciencias Na-
turales, Buenos Aires, Argentina.
Anatomical
ang
art
bocc
d
ect
exoc
fit
fm
fr
ju
la
m
n
P
pm
Pi
po
PP
P rf
pt
q
qj
rart
soc
spl
sq
sur
angular
articular
basioccipital
dentary
ectopterygoid
exoccipital
foramen intertympanicum
foramen magnum
frontal
jugal
lacrymal
maxilla
nasal
parietal
premaxilla
palatine
postorbital
posterior palpebral
prefrontal
pterygoid
quadrate
quadratojugal
retroarticular process
supraoccipital
splenial
squamosal
surangular
Character definitions 1-101 were taken
from Clark (1994) and have the same numer¬
ation as in the original publication. Character
5 was excluded from the analysis (because it
depends on the modified definition of charac¬
ter 6); however, its inclusion does not affect
the outcome of the analysis (except for the tree
length). The additional characters are also
listed here and their respective source is cited
along with the character number of the
original publication. Characters 1, 3, 6, 23,
37, 45, 49, 65, 67, 69, 73, 77, 79, 90, 91, 96, 97,
103, 104, 105, 107, 126, 143, 149, and 165 were
set as ordered characters (marked “+” in this
list).
Character 1 (modified from Clark, 1994;
character 1). + External surface of dorsal
cranial bones: smooth (0), slightly grooved (1),
and heavily ornamented with deep pits and
grooves (2).
Character 2 (modified from Clark, 1994;
character 2). Skull expansion at orbits: grad¬
ual (0) or abrupt (1).
Character 3 (modified from Clark, 1994;
character 3). + Rostrum proportions: narrow
oreinirostral (0) or broad oreinirostral (1) or
nearly tubular (2) or platyrostral (3).
Character 4 (Clark, 1994; character 4).
Premaxilla participation in internarial bar:
forming at least the ventral half (0) or with
little participation (1).
Character 5 (Clark, 1994; character 5).
Premaxilla anterior to nares: narrow (0) or
broad (1).
Character 6 (modified from Clark, 1994;
character 6). + External nares facing: ante-
rolaterally or anteriorly (0), dorsally not
separated by premaxillary bar from anterior
2006
ZAHER ET AL.: CRANIAL MORPHOLOGY OF MARILIASUCHUS AMARALI
29
edge of rostrum (1), or dorsally separated by
premaxillary bar (2).
Character 7 (Clark, 1994; character 7).
Palatal parts of premaxillae: do not meet
posterior to incisive foramen (0) or meet
posteriorly along contact with maxillae (1).
Character 8 (Clark, 1994; character 8).
Premaxilla-maxilla contact: premaxilla loose¬
ly overlies maxilla (0) or sutured together
along a butt joint (1).
Character 9 (modified from Clark, 1994;
character 9). Ventrally opened notch on
ventral edge of rostrum at premaxilla-maxilla
contact: absent (0) or present as a notch (1) or
present as a large fenestra (2).
Character 10 (Clark, 1994; character 10).
Posterior ends of palatal branches of maxillae
anterior to palatines: do not meet (0) or meet
( 1 ).
Character 11 (Clark, 1994; character 11).
Nasal-lacrimal contact: (0) or do not contact
( 1 ).
Character 12 (Clark, 1994; character 12).
Lacrimal contacts nasal along: medial edge
only (0) or medial and anterior edges (1).
Character 13 (Clark, 1994; character 13).
Nasal contribution to narial border: yes (0) or
no (1).
Character 14 (Clark, 1994; character 14).
Nasal-premaxilla contact: present (0) or ab¬
sent (1).
Character 15 (modified from Clark, 1994;
character 15). Descending process of prefron¬
tal: does not contact palate (0) or contacts
palate (1).
Character 16 (Clark, 1994; character 16).
Postorbital-jugal contact: postorbital anterior
to jugal or postorbital medial to jugal (1) or
postorbital lateral to jugal (2).
Character 17 (Clark, 1994; character 17).
Anterior part of the jugal with respect to
posterior part: as broad (0) or twice as broad (1).
Character 18 (Clark, 1994; character 18).
Jugal bar beneath infratemporal fenestra:
flattened (0) or rod-shaped (1).
Character 19 (Clark, 1994; character 19).
Quadratojugal dorsal process: narrow, con¬
tacting only a small part of postorbital (0) or
broad, extensively contacting the postorbital
( 1 ).
Character 20 (Clark, 1994; character 20).
Frontal width between orbits: narrow, as
broad as nasals (0) or broad, twice as broad
as nasals (1).
Character 21 (Clark, 1994; character 21).
Frontals: paired (0), unpaired (1).
Character 22 (Clark, 1994; character 22).
Dorsal surface of frontal and parietal: flat (0)
or with midline ridge (1).
Character 23 (modified from Clark, 1994;
character 23; by Buckley and Brochu, 1999;
character 81). + Parieto-postorbital suture:
absent from dorsal surface of skull roof and
supratemporal fossa (0) or absent from dorsal
surface of skull roof but broadly present
within supratemporal fossa (1) or present
within supratemporal fossa and on dorsal
surface of skull roof (2).
Character 24 (Clark, 1994; character 24).
Supratemporal roof dorsal surface: complex
(0) or dorsally flat “skull table” developed,
with postorbital and squamosal with flat
shelves extending laterally beyond quadrate
contact (1).
Character 25 (modified from Clark, 1994;
character 25). Postorbital bar: sculpted (if
skull sculpted) (0) or unsculpted (1).
Character 26 (modified from Clark, 1994;
character 26). Postorbital bar: transversely
flattened (0) or cylindrical (1).
Character 27 (Clark, 1994; character 27).
Vascular opening in dorsal surface of post¬
orbital bar: absent (0), present (1).
Character 28 (modified from Clark, 1994;
character 28). Postorbital anterolateral pro¬
cess: absent or poorly developed (0) or well
developed, long, and acute (1).
Character 29 (Clark, 1994; character 29).
Dorsal part of the postorbital: with anterior
and lateral edges only (0) or with anterolat-
erally facing edge (1).
Character 30 (Clark, 1994; character 30).
Dorsal end of the postorbital bar broadens
dorsally, continuous with dorsal part of post¬
orbital (0) or dorsal part of the postorbital bar
constricted, distinct from the dorsal part of the
postorbital (1).
Character 31 (Clark, 1994; character 31).
Bar between orbit and supratemporal fossa
broad and solid, with broadly sculpted dorsal
surface (0) or bar narrow, sculpting restricted
to anterior surface (1).
Character 32 (modified from Clark, 1994;
character 32). Parietal: with broad occipital
30
AMERICAN MUSEUM NOVITATES
NO. 3512
portion (0) or without broad occipital portion
( 1 ).
Character 33 (Clark, 1994; character 33).
Parietal: with broad sculpted region separat¬
ing fossae (0) or with sagittal crest between
supratemporal fossae (1).
Character 34 (Clark, 1994; character 34).
Postparietal (dermosupraoccipital): a distinct
element (0) or not distinct (fused with parie¬
tal?) (1).
Character 35 (Clark, 1994; character 35).
Posterodorsal corner of the squamosal:
squared off, lacking extra “lobe” (0) or with
unsculptured “lobe” (1).
Character 36 (modified from Clark, 1994;
character 36). Posterolateral process of squa¬
mosal: poorly developed and projected hor¬
izontally at the same level of the skull (0) or
elongated, thin, and posteriorly directed, not
ventrally deflected (1) or elongated, poster-
olaterally directed, and ventrally deflected
( 2 ).
Character 37. (Clark, 1994; character 37). +
Palatines: do not meet on palate below the
narial passage (0) or form palatal shelves that
do not meet (1) or meet ventrally to the narial
passage, forming part of secondary palate (2).
Character 38 (Clark, 1994; character 38).
Pterygoid: restricted to palate and suspensor-
ium, joints with quadrate and basisphenoid
overlapping (0) or pterygoid extends dorsally
to contact laterosphenoid and form ventrolat¬
eral edge of the trigeminal foramen, strongly
sutured to quadrate and laterosphenoid (1).
Character 39 (modified from Clark, 1994;
character 39). Choanal opening: continuous
with pterygoid ventral surface except for
anterior and anterolateral borders (0) or opens
into palate through a deep midline depression
(choanal groove) (1).
Character 40 (Clark, 1994; character 40).
Palatal surface of pterygoids: smooth (0) or
sculpted (1).
Character 41 (Clark, 1994; character 41).
Pterygoids posterior to choanae: separated (0)
or fused (1).
Character 42 (modified from Clark, 1994;
character 42; by Ortega et al., 2000; character
139). Depression on primary pterygoidean
palate posterior to choana: absent or moder¬
ate in size being narrower than palatine bar (0)
or wider than palatine bar (1).
Character 43 (Clark, 1994; character 43).
Pterygoids: do not enclose choana (0) or
enclose choana (1).
Character 44 (modified from Clark, 1994;
character 44). Anterior edge of choanae
situated near posterior edge of suborbital
fenestra (or anteriorly) (0) or near posterior
edge of pterygoid flanges (1).
Character 45 (Clark, 1994; character 45). +
Quadrate: without fenestrae (0) or with single
fenestrae (1) or with three or more fenestrae
on dorsal and posteromedial surfaces (2).
Character 46 (Clark, 1994; character 46).
Posterior edge of quadrate: broad medial to
tympanum, gently concave (0) or posterior
edge narrow dorsal to otoccipital contact,
strongly concave (1).
Character 47 (Clark, 1994; character 47).
Dorsal, primary head of quadrate articulates
with: squamosal, otoccipital and prootic (0) or
with prootic and laterosphenoid (1).
Character 48 (Clark, 1994; character 48).
Ventrolateral contact of otoccipital with
quadrate: very narrow (0) or broad (1).
Character 49 (Clark, 1994; character 49). +
Quadrate, squamosal, and otoccipital: do not
meet to enclose cranioquadrate passage (0) or
enclose passage near lateral edge of skull (1) or
meet broadly lateral to the passage (2).
Character 50 (Clark, 1994; character 50).
Pterygoid ramus of quadrate: with flat ventral
edge (0) or with deep groove along ventral
edge (1).
Character 51 (Clark, 1994; character 51).
Ventromedial part of quadrate: does not
contact otoccipital (0) or contacts otoccipital
to enclose carotid artery and form passage for
cranial nerves IX-XI (1).
Character 52 (Clark, 1994; character 52).
Eustachian tubes: not enclosed between ba-
sioccipital and basisphenoid (0) or entirely
enclosed (1).
Character 53 (Clark, 1994; character 53).
Basisphenoid rostrum (cultriform process):
slender (0) or dorsoventrally expanded (1).
Character 54 (Clark, 1994; character 54).
Basipterygoid process: prominent, forming
movable joint with pterygoid (0) or basipter¬
ygoid process small or absent, with basi¬
sphenoid joint suturally closed (1).
Character 55 (modified from Clark, 1994;
character 55; by Ortega et al., 2000; character
2006
ZAHER ET AL.: CRANIAL MORPHOLOGY OF MARILIASUCHUS AMARALI
31
68). Basisphenoid ventral surface: shorter than
the basioccipital (0) or wide and similar to or
longer in length than basioccipital (1).
Character 56 (Clark, 1994; character 56).
Basisphenoid: exposed on ventral surface of
braincase (0) or virtually excluded from
ventral surface by pterygoid and basioccipital
( 1 ).
Character 57 (Clark, 1994; character 57).
Basioccipital: without well-developed biltaeral
tuberosities (0) or with large pendulous tubera
( 1 ).
Character 58 (Clark, 1994; character 58).
Otoccipital: without laterally concave des¬
cending flange ventral to subcapsular process
(0) or with flange (1).
Character 59 (Clark, 1994; character 59).
Cranial nerves IX-XI: pass through common
large foramen vagi in otoccipital (0) or cranial
nerve IX pass medial to nerves X and XI in
separate passage (1).
Character 60 (Clark, 1994; character 60).
Otoccipital: without large ventrolateral part
ventral to paroccipital process (0) or with
large ventrolateral part (1).
Character 61 (Clark, 1994; character 61).
Crista interfenestralis between fenestrae pseu¬
dorotunda and ovalis nearly vertical (0) or
horizontal (1).
Character 62 (Clark, 1994; character 62).
Supraoccipital: forms dorsal edge of the
foramen magnum (0) or otoccipitals broadly
meet dorsal to the foramen magnum, separat¬
ing supraoccipital from foramen (1).
Character 63 (Clark, 1994; character 63).
Mastoid antrum: does not extend into su¬
praoccipital (0) or extends through transverse
canal in supraoccipital to connect middle ear
regions (1).
Character 64 (Clark, 1994; character 64).
Posterior surface of supraoccipital: nearly flat
(0), or with bilateral posterior prominences
( 1 ).
Character 65 (modified from Clark, 1994;
character 65). + One small palpebral present in
orbit (0) or one large palpebral (1) or two large
palpebrals (2).
Character 66 (Clark, 1994; character 66).
External nares: divided by a septum (0) or
confluent (1).
Character 67 (Clark, 1994; character 67). +
Antorbital fenestra: as large as orbit (0) or
about half the diameter of the orbit (1) or
much smaller than the orbit (2) or absent (3).
Character 68 (modified from Clark, 1994;
character 68; by Ortega et al., 2000; character
41). Supratemporal fenestrae extension: rela¬
tively large, covering most of surface of
skull roof (0) or relatively short, fenestrae
surrounded by a flat and extended skull roof
( 1 ).
Character 69 (modified from Clark, 1994;
character 69). + Choanal groove: undivided
(0) or partially septated (1) or completely
septated (2).
Character 70 (Clark, 1994; character 70).
Dentary: extends posteriorly beneath mandib¬
ular fenestra (0) or does not extend beneath
fenestra (1).
Character 71 (modified from Clark, 1994;
character 71). Retroarticular process: absent
or extremely reduced (0) or very short, broad,
and robust (1) or with an extensive rounded,
wide, and flat (or slightly concave) surface
projected posteroventrally and facing dor-
somedially (2) or posteriorly elongated, tri¬
angular shaped and facing dorsally (3) or
posteroventrally projecting and paddle shaped
(4).
Character 72 (Clark, 1994; character 72).
Prearticular: present (0) or absent (1).
Character 73 (modified from Clark, 1994;
character 73). + Articular without medial
process (0) or with short process not contact¬
ing braincase (1) or with process articulating
with otoccipital and basisphenoid (2).
Character 74 (Clark, 1994; character 74).
Dorsal edge of surangular: flat (0) or arched
dorsally (1).
Character 75 (Clark, 1994; character 75).
Mandibular fenestra: present (0) or absent (1).
Character 76 (Clark, 1994; character 76).
Insertion area for M. pterygoideous posterior:
does not extend onto lateral surface of angular
(0) or extends onto lateral surface of angular
( 1 ).
Character 77 (modified from Clark, 1994;
character 77). + Splenial involvement in
symphysis in ventral view: not involved (0)
or involved slightly in symphysis (1) or
extensively involved (2).
Character 78 (Clark, 1994; character 78).
Posterior premaxillary teeth: similar in size to
anterior teeth (0) or much longer (1).
32
AMERICAN MUSEUM NOVITATES
NO. 3512
Character 79 (modified from Clark, 1994;
character 79). + Maxillary teeth waves: absent,
no tooth size variation (0) or one wave of teeth
enlarged (1) or enlarged maxillary teeth curved
in two waves (“festooned”) (2).
Character 80 (Clark, 1994; character 80).
Anterior dentary teeth opposite premaxilla-
maxilla contact: no more than twice the length
of other dentary teeth (0) or more than twice
the length (1).
Character 81 (modified from Clark, 1994;
character 81). Dentary teeth posterior to tooth
opposite premaxilla-maxilla contact: equal in
size (0) or enlarged dentary teeth opposite to
smaller teeth in maxillary tooth row (1).
Character 82 (modified from Clark, 1994;
character 82; by Ortega et al., 2000; character
120) . Anterior and posterior scapular edges:
symmetrical in lateral view (0) or anterior
edge more strongly concave than posterior
edge (1) or dor sally narrow with straight edges
( 2 ).
Character 83 (modified from Clark, 1994;
character 83; by Ortega et al., 2000; character
121) . Coracoid length: up to two-thirds of the
scapular length (0) or subequal in length to
scapula (1).
Character 84 (Clark, 1994; character 84).
Anterior process of ilium: similar in length to
posterior process (0) or one-quarter or less of
the length of the posterior process (1).
Character 85 (Clark, 1994; character 85).
Pubis: rodlike without expanded distal end (0)
or with expanded distal end (1).
Character 86 (Clark, 1994; character 86).
Pubis: forms anterior half of ventral edge of
acetabulum (0) or pubis at least partially
excluded from the acetabulum by the anterior
process of the ischium (1).
Character 87 (Clark, 1994; character 87).
Distal end of femur: with large lateral facet for
the fibula (0) or with very small facet (1).
Character 88 (Clark, 1994; character 88).
Fifth pedal digit: with phalanges (0) or
without phalanges (1).
Character 89 (Clark, 1994; character 89).
Atlas intercentrum: broader than long (0) or
as long as broad (1).
Character 90 (modified from Clark, 1994;
character 90). + Cervical neural spines: all
anteroposteriorly large (0), only posterior ones
rodlike (1), or all spines rodlike (2).
Character 91 (modified from Clark, 1994;
character 91; by Buscalioni and Sanz, 1988;
and by Brochu, 1997; character 37 and
character 7, respectively). + Hypapophyses in
cervicodorsal vertebrae: absent (0) or present
only in cervical vertebrae (1) or present in
cervical and the first two dorsal vertebrae (2)
or present up to the third dorsal vertebra (3)
or up to the fourth dorsal vertebrae (4).
Character 92 (Clark, 1994; character 92).
Cervical vertebrae: amphicoelous or amphy-
platian (0) or procoelous (1).
Character 93 (Clark, 1994; character 93).
Trunk vertebrae: amphicoelous or amphypla-
tian (0) or procoelous (1).
Character 94 (Clark, 1994; character 94).
All caudal vertebrae: amphicoelous or am-
phyplatian (0) or first caudal biconvex with
other procoelous (1) or procoelous (2).
Character 95 (Clark, 1994; character 95).
Dorsal osteoderms: rounded or ovate (0) or
rectangular, broader than long (1) or square (2).
Character 96 (modified from Clark, 1994;
character 96; and Brochu, 1997; character 40).
+ Dorsal osteoderms without articular anteri¬
or process (0) or with a discrete convexity on
anterior margin (1) or with a well-developed
process located anterolaterally in dorsal para¬
sagittal osteoderms (2).
Character 97 (modified from Clark, 1994;
character 97; by Ortega et al., 2000; characters
107 and 108). + Rows of dorsal osteoderms:
two parallel rows (0) or more than two (1) or
more than four with “accessory ranges of
osteoderms” (sensu Frey, 1988) (2).
Character 98 (Clark, 1994; character 98).
Osteoderms: Some or all imbricated (0) or
sutured to one another (1).
Character 99 (Clark, 1994; character 99).
Tail osteoderms: dorsal only (0) or completely
surrounded by osteoderms (1).
Character 100 (Clark, 1994; character 100).
Trunk osteoderms: absent from ventral part of
the trunk (0) or present (1).
Character 101 (Clark, 1994; character 101).
Osteoderms: with longitudinal keels on dorsal
surfaces (0) or without longitudinal keels (1).
Character 102 (Wu and Sues, 1996; charac¬
ter 14). Jugal: participating in margin of
antorbital fossa (0) or separated from it (1).
Character 103 (modified from Wu and Sues,
1996; character 23). + Articular facet for
2006
ZAHER ET AL.: CRANIAL MORPHOLOGY OF MARILIASUCHUS AMARALI
33
quadrate condyle: equal in length to the
quadrate condyles (0) or slightly longer (1)
or close to three times the length of the
quadrate condyles (2).
Character 104 (modified from Wu and Sues,
1996, and Wu et al., 1997; character 24 and
character 124, respectively). + Jaw joint:
placed at level with basioccipital condyle (0)
or below basioccipital condyle about above
level of lower tooth row (1) or below level of
tooth row (2).
Character 105 (modified from Wu and Sues,
1996, and Ortega et al., 2000; character 27 and
character 133, respectively). + Premaxillary
teeth: five (0), four (1), three (2), or two (3).
Character 106 (modified from Wu and Sues,
1996; character 29). Unsculptured region
along alveolar margin on lateral surface of
maxilla: absent (0) or present (1).
Character 107 (modified from Wu and Sues,
1996; character 30). + Maxilla: with eight or
more teeth (0) or seven (1) or six (2) or five (3)
or four (4) teeth.
Character 108 (Wu and Sues, 1996; charac¬
ter 33). Coracoid: without posteromedial or
ventromedial process (0) or with elongate
posteromedial process (1) or distally expanded
ventromedial process (2).
Character 109 (Wu and Sues, 1996; charac¬
ter 40). Radiale and ulnare: short and massive
(0) or elongate (1).
Character 110 (Wu and Sues, 1996; charac¬
ter 41). Postacetabular process: directed pos-
teroventrally or posteriorly (0) or directed
posterodorsally and much higher in position
than preacetabular process (1).
Character 111 (modified from Gomani,
1997; character 4). Prefrontals anterior to
orbits: elongated, oriented parallel to ante¬
roposterior axis of the skull (0) or short and
broad, oriented posteromedially-anterolater-
ally (1).
Character 112 (modified from Gomani,
1997; character 32). Basioccipital and ventral
part of otoccipital: facing posteriorly (0) or
posteroventrally (1).
Character 113 (Buscalioni and Sanz, 1988;
character 35). Vertebral centra: cylindrical (0)
or spool shaped (1).
Character 114 (modified from Buscalioni
and Sanz, 1988; character 39). Transverse
process of posterior dorsal vertebrae dorso-
ventrally low and laminar (0) or dorsoventral-
ly high (1).
Character 115 (Buscalioni and Sanz, 1988;
character 44). Number of sacral vertebrae: two
(0) or more than two (1).
Character 116 (Buscalioni and Sanz, 1988;
character 49). Supra-acetabular crest: present
(0) or absent (1).
Character 117 (Buscalioni and Sanz, 1988;
character 54). Proximal end of radiale ex¬
panded symmetrically, similarly to the distal
end (0) or more expanded proximomedially
than proximolaterally (1).
Character 118 (Ortega et al., 1996; character
5). Lateral surface of the dentary: without
a longitudinal depression (0) or with a longi¬
tudinal depression (1).
Character 119 (Ortega et al., 1996; character
9). Ventral exposure of splenials: absent (0) or
present (1).
Character 120 (Ortega et al., 1996, 2000;
characters 11 and 100, respectively). Tooth
margins: with denticulate carinae (0) or
without carinae or with smooth or crenulated
carinae (1).
Character 121 (modified from Pol, 1999a,
and Ortega et al., 2000; character 133 and
character 145, respectively). Lateral surface of
anterior process of jugal: flat or convex (0) or
with broad shelf below the orbit with tri¬
angular depression underneath it (1).
Character 122 (Pol, 1999a; character 134).
Jugal: does not exceed the anterior margin of
orbit (0) or exceeds (1).
Character 123 (Pol, 1999a; character 135).
Notch in premaxilla on lateral edge of external
nares: absent (0) or present on the dorsal half
of the external nares lateral margin (1).
Character 124 (Pol, 1999a; character 136).
Dorsal border of external nares: formed
mostly by the nasals (0) or by both the nasals
and premaxilla (1).
Character 125 (Pol, 1999a; character 138).
Posterodorsal process of premaxilla: absent
(0) or present extending posteriorly wedging
between maxilla and nasals (1).
Character 126 (Pol, 1999a, and Ortega et
al., 2000; character 139 and character 9,
respectively). + Premaxilla-maxilla suture
in palatal view, medial to alveolar region:
anteromedially directed (0) or sinusoidal,
posteromedially directed on its lateral
34
AMERICAN MUSEUM NOVITATES
NO. 3512
half and anteromedially directed along its
medial region (1) or posteromedially directed
( 2 ).
Character 127 (Pol, 1999a; character 140).
Nasal lateral border posterior to external
nares: laterally concave (0) or straight (1).
Character 128 (Pol, 1999a; character 141).
Nasal lateral edges: nearly parallel (0) or
entirely oblique to each other converging
anteriorly (1) or oblique to each other di¬
verging anteriorly (2).
Character 129 (Pol, 1999a; character 143).
Palatine anteromedial margin: exceeding the
anterior margin of the palatal fenestrae
wedging between the maxillae (0) or not
exceeding the anterior margin of palatal
fenestrae (1).
Character 130 (Pol, 1999a; character 144).
Dorsoventral height of jugal antorbital region
respect to infraorbital region: equal or lower
(0) or antorbital region more expanded than
infraorbital region of jugal (1).
Character 131 (Pol, 1999a; character 145).
Maxilla-lacrimal contact: partially included
in antorbital fossa (0) or completely included
( 1 ).
Character 132 (Pol, 1999a; character 146).
Lateral eustachian tube openings: located
posteriorly to the medial opening (0) or
aligned anteroposteriorly and dorsoventrally
( 1 ).
Character 133 (Pol, 1999a; character 147).
Anterior process of ectopterygoid: developed
(0) or reduced-absent (1).
Character 134 (Pol, 1999a; character 148).
Posterior process of ectopterygoid: developed
(0) or reduced-absent (1).
Character 135 (Pol, 1999a, and Ortega et
al., 2000; character 149 and character 13,
respectively). Small foramen located in the
premaxillo-maxillary suture in lateral surface
(not for big mandibular teeth): absent (0) or
present (1).
Character 136 (Pol, 1999a; character 150).
Jugal posterior process: exceeding pos¬
teriorly the infratemporal fenestrae (0) or not
( 1 ).
Character 137 (Pol, 1999a; character 151).
Compressed crown of maxillary teeth: orient¬
ed parallel to the longitudinal axis of skull (0)
or obliquely disposed (1).
Character 138 (Pol, 1999a; character 152).
Large and aligned neurovascular foramina on
lateral maxilary surface: absent (0) or present
( 1 ).
Character 139 (modified from Pol, 1999a;
character 153). External surface of maxilla
and premaxilla: with a single plane facing
laterally (0) or with ventral region facing
laterally and dorsal region facing dorsolater-
ally (1).
Character 140 (Pol, 1999a, and Ortega et
al., 2000; character 154 and character 104,
respectively). Maxillary teeth: not compressed
laterally (0) or compressed laterally (1).
Character 141 (Pol, 1999a; character 155).
Posteroventral corner of quadratojugal: reach¬
ing the quadrate condyles (0) or not reaching
the quadrate condyles (1).
Character 142 (Pol, 1999a; character 156).
Base of postorbital process of jugal: directed
posterodorsally (0) or dorsally (1).
Character 143 (Pol, 1999a; character 157). +
Postorbital process of jugal: anteriorly placed
(0), in the middle (1), or posteriorly positioned
( 2 ).
Character 144 (Pol, 1999a, and Ortega et
al., 2000; character 158 and character 36,
respectively). Postorbital-ectopterygoid con¬
tact: present (0), absent (1).
Character 145 (Pol, 1999a; character 161).
Quadratojugal: not ornamented (0) or orna¬
mented in the base (1).
Character 146 (Pol, 1999a; character 162).
Prefrontal-maxillary contact in the inner
anteromedial region of orbit: absent (0) or
present (1).
Character 147 (Pol, 1999a; character 163).
Basisphenoid: without lateral exposure (0) or
with lateral exposure on the braincase (1).
Character 148 (Pol, 1999a; character 165).
Quadrate process of pterygoids: well devel¬
oped (0) or poorly developed (1).
Character 149 (modified from Pol, 1999a,
and Ortega et al., 2000; character 166 and
character 44, respectively). + Quadrate major
axis directed: posteroventrally (0) or ventrally
(1) or anteroventrally (2).
Character 150 (Pol, 1999a; character 167).
Quadrate distal end: with only one plane
facing posteriorly (0) or with two distinct
faces in posterior view, a posterior one and
2006
ZAHER ET AL.: CRANIAL MORPHOLOGY OF MARILIASUCHUS AMARALI
35
a medial one bearing the foramen aereum
( 1 ).
Character 151 (Pol, 1999a; character 168).
Anteroposterior development of neural spine
in axis: well developed, covering all the neural
arch length (0) or poorly developed, located
over the posterior half of the neural arch
( 1 ).
Character 152 (Pol, 1999a; character 169).
Prezygapophyses of axis: not exceeding ante¬
rior edge of neural arch (0) or exceeding the
anterior margin of neural arch (1).
Character 153 (Pol, 1999a; character 170).
Postzygapophyses of axis: well developed,
curved laterally (0) or poorly developed
( 1 ).
Character 154 (modified from Pol, 1999b;
character 212). Shape of dentary symphysis in
ventral view: tapering anteriorly forming an
angle (0) or U shaped, smoothly curving
anteriorly (1) or lateral edges longitudinally
oriented, convex anterolateral corner, and
extensive transversally oriented anterior edge
( 2 ).
Character 155 (Pol, 1999b; character 213).
Unsculpted region in the dentary below the
tooth row: absent (0) or present (1).
Character 156 (Ortega et al., 1996, and
Buckley et al., 2000; character 13). Cheek
teeth: not constricted at base of crown (0) or
constricted (1).
Character 157 (Ortega et al., 2000; character
42). Outer surface of squamosal laterodorsally
oriented: extensive (0) or reduced and sculpted
(1) or reduced and unsculpted (2).
Character 158 (Ortega et al., 2000; character
74). Length/height proportion of infratempor¬
al fenestra: higher than wide or equal (0) or
very antero-posteriorly elongated (1).
Character 159 (Ortega et al., 2000; character
90). Foramen intramandibularis oralis: small
or absent (0) or big and slotlike (1).
Character 160 (Ortega et al., 2000; character
146). Ectopterygoid medial process: single (0)
or forked (1).
Character 161 (modified from Gomani,
1997, and Buckley et al., 2000; character 46
and character 113, respectively). Cusps of
teeth: unique cusp (0), one main cusp with
smaller cusps arranged in one row (1) or one
main cusp with smaller cusps arranged in
more than one row (2) or several cusps of
equal size arranged in more than one row (3)
or multiple small cusps along edges of
occlussal surface (4).
Character 162 (Pol and Norell, 2004a;
character 164). Cross section of distal end
of quadrate: mediolaterally wide and ante-
roposteriorly thin (0) or subquadrangular
( 1 ).
Character 163 (Pol and Norell, 2004a;
character 165). Palatine-pterygoid contact on
palate: palatines overlie pterygoids (0) or
palatines firmly sutured to pterygoids (1).
Character 164 (Wu et al., 1997; character
103). Squamosal descending process: absent
(0) or present (1).
Character 165 (modified from Wu et al.,
1997; character 105). + Development of distal
quadrate body ventral to otoccipital-quadrate
contact: distinct (0) or incipiently distinct (1)
or indistinct (2).
Character 166 (Wu et al., 1997; character
106). Pterygoid flanges: thin and laminar (0)
or dorsoventrally thick, with pneumatic spaces
( 1 ).
Character 167 (Wu et al., 1997; character
108) . Postorbital participation in infratempor¬
al fenestra: almost or entirely excluded (0) or
bordering infratemporal fenestra (1).
Character 168 (Wu et al., 1997; character
109) . Palatines: form margin of suborbital
fenestra (0) or excluded from margin of
suborbital fenestra (1).
Character 169 (Wu et al., 1997; character
110) . Angular posterior to mandibular fenes¬
tra: widely exposed on lateral surface of
mandible (0) or shifted to the ventral surface
of mandible (1).
Character 170 (Wu et al., 1997; character
112). Posteroventral edge of mandibular
ramus: straight or convex (0) or markedly
deflected (1).
Character 171 (modified from Wu et al.,
1997; character 119). Quadrate ramus of
pterygoid in ventral view: narrow (0) or broad
( 1 ).
Character 172 (Wu et al., 1997; character
121) . Pterygoids: not in contact anterior to
basisphenoid on palate (0) or pterygoids in
contact (1).
Character 173 (Wu et al., 1997; character
122) . Olecranon: well developed (0) or absent
( 1 ).
36
AMERICAN MUSEUM NOVITATES
NO. 3512
Character 174 (Wu et al., 1997; character
123). Cranial table width respect to ventral
portion of skull: as wide as ventral portion (0)
or narrower than ventral portion of skull (1).
Character 175 (Wu et al., 1997; character
127) . Depression on posterolateral surface of
maxilla: absent (0) or present (1).
Character 176 (Wu et al., 1997; character
128) . Anterior palatal fenestra: absent (0) or
present (1).
Character 177 (Pol and Norell, 2004a;
character 179). Paired ridges located medially
on ventral surface of basisphenoid: absent (0)
or present (1).
Character 178 (Pol and Norell, 2004a;
character 180). Posterolateral end of quad-
ratojugal: acute or rounded, tightly overlap¬
ping the quadrate (0) or with sinusoidal
ventral edge and wide and rounded posterior
edge slightly overhanging the lateral surface of
the quadrate (1).
Character 179 (Pol and Norell, 2004a;
character 181). Orientation of quadrate body
distal to otoccipital-quadrate contact in pos¬
terior view: ventrally (0) or ventrolaterally (1).
Character 180 (Gasparini et al., 1991;
character 3). Wedgelike process of the maxilla
in lateral surface of premaxilla-maxilla suture:
absent (0) or present (1).
Character 181 (Pol and Norell, 2004b;
character 181). Palpebrals: separated from
the lateral edge of the frontals (0) or fused to
each other and the lateral margin of the
frontals (1).
Character 182 (Pol and Norell, 2004b;
character 182). External surface of ascending
process of jugal: exposed laterally (0) or
exposed posterolaterally (1).
Character 183 (Pol and Norell, 2004b;
character 183). Longitudinal ridge on lateral
surface of jugal below infratemporal fenestra:
absent (0) or present (1).
Character 184 (Pol and Norell, 2004b;
character 184). Dorsal surface of posterolat¬
eral region of squamosal: without ridges (0) or
with three curved ridges oriented longitudi¬
nally (1).
Character 185 (Pol and Norell, 2004b;
character 185). Ridge along dorsal section of
quadrate-quadratojugal contact: absent (0) or
present (1).
Character 186 (Pol and Norell, 2004b;
character 186). Sharp ridge along the ventral
surface of angular: absent (0) or present (1).
Character 187 (Pol and Norell, 2004b;
character 187). Longitudinal ridge along the
dorsolateral surface of surangular: absent (0)
or present (1).
Character 188 (Pol and Norell, 2004b;
character 188). Dorsal surface of osteoderms
ornamented with anterolaterally and antero-
medially directed ridges (fleur de lys pattern of
Osmolska et al., 1997): absent (0) or present
( 1 ).
Character 189 (Pol and Norell, 2004b;
character 189). Cervical region surrounded
by lateral and ventral osteoderms sutured to
the dorsal elements: absent (0) or present (1).
Character 190 (Pol and Norell, 2004b;
character 190). Appendicular osteoderms:
absent (0) or present (1).
Character 191 (Ortega et al., 2000; character
72). Supratemporal fenestra: present (0) ab¬
sent (1).
Character 192. Ventral half of lacrimal:
extending posteroventrally, widely contacting
the jugal (0) or tapers posteroventrally, not
contacting or contacting slightly the jugal (1).
Character 193. Large foramen on lateral
surface of anterior jugal: absent (0) or present
( 1 ).
Character 194. Procumbent premaxilary
and anterior dentary alveoli: absent (0) or
present (1).
Character 195. Palatine bar: absent (0) or
present (1).
Character 196. Participation of ectoptery-
goid on palatine bar: no (0) or yes (1).
Character 197 (Pol and Norell, 2004b;
character 192). Choanal opening: opened
posteriorly and continuous with pterygoid
surface (0) or closed posteriorly by an elevated
wall formed by the pterygoids (1).
Character 198. Ectopterygoid: projecting
medially on ventral surface of pterygoid
flanges barely extended (0) or widely extended,
covering approximately the lateral half of the
ventral surface of the pterygoid flanges (1).
2006
ZAHER ET AL.: CRANIAL MORPHOLOGY OF MARILIASUCHUS AMARALI
37
APPENDIX 3
Data Matrix Used in Phylogenetic Analysis
The data matrix contains 46 taxa and
198 characters.
010??0????00??0?1200101101112?????0????011-
00700701000111?101001?01?10000000011?0??-
40012??00011??0?00???00?????????????0?
Zaraasuchus
Gracilisuchus
000000 ?? 0 ? 000000000000 ? 0 ? 000000000 ? 0 ?? 0 ?-
0 ? 00000 ? 000 ??? 0000 ? 0 ??? 00000 ? 100000 ? 00000 -
000?0???0000?0?000001012?00?00????0?017010-
00?? 1 ?01 ???000001002?0???0000?????0??0??000-
0?00000?0?00000000?0?0000??0??0?
10????????????? 1 ?01 ?01 ?1000001? 10?02?????????-
???????????????????2????? 1 ??010?????????????? {12-
34} 0?? 1010??0???????????0??????????????????????-
0????? 1 ?? 1 ????????? 1 ?00???????? 1 ?00???0?????? 1 -
1111111111???????
Gobiosuchus
Terrestrisuchus
000 ?? 00 ?? 0 ?? 000000 ? 000 ? 0 ? 00 ? 000 ? 110 ? 00000 -
? 00000 ? 000 ?? 0 ? 000 ? 000 ???? 00 ??? 010 ?? 0 ? 00000 -
0?010?0000?0200000101??01100??00000?0010-
0 ?? 10 ? 00 ? 110 ? 0 ? 0 ??( 01 ) 110 ??? 00000 ???? 00 ? 0 ??-
0 ???? 00 ?? 0 ? 0 ??? 0 ??????????? 0 ????? 0 ?
Dibothrosuchus
000?00?020??001 ???000000??????00110000000?-
00000 ? 0000 ? 00000 ? 0?0101000?010100?0010 ? 0 -
00?????2000?0?????01010701100?0?0000000100-
1 ? 10?00?17000101011100??000001 ??00000010-
001000100?0?00000000?000000?00??0?
Protosuchus
1010007110000011001 ?(01 )(01)? 1700001 ? 10?0-
201000?0020112011111000?0????201 ??? 172010-
0(01)010?0? 1 ???????0? 1010110{01}012002??00-
00???0010(01 )00001000000?000010012117000-
0???110000000?121000011?00?0?00111111111-
11?00??00
Sichuanosuchus
(12)01 ??0?1200(01)00? 10010(01)17110??? 1 ?00?-
021 ? 10?00020? 1 ?011?1100???????2?11????1?00-
0011 ? 1 ?? 1 ?????000???????? 1 ? 11 ?0? 1 ????0?? 1001-
00?? 1 ?? 10?0????00110(01)1210??00????? 1 ?????0-
101110111111007110000100?1???0?00??00
Shantungosuchus
2100000120?0000110100021000001000100010-
107002010011111100101011020117110210001 -
010100011100 { 1234 } 00 ? 120011010111021001 -
010000{01}000000?01??01??10010{01}010100-
0000???0100000000120000011110??01000?010-
?0000?00??0?
Hemiprotosuchus
? 00 ? 00 ? 10 ?????? 10010 ? 0 ?? 00?0010 ? 11 ? 0??01 ?? 0 -
020?00? 1171100101 ?? 1 ?2? 11 ?? 1 ?21 ????01 ??????-
???0???? 1200? 1 ? 101 ??0??????????000?000?? 10?0-
0???00000?? 10?????00???????0?000? 12???001 ?? 1 -
0 ? 0 ? 00 ? 01 ?? 00 ??? 0 ????? 0 ?
Orthosuchus
21100001201?0001001000{01}1000001000100-
0?000?002011001111100??1?1?02011?0?0?0?00-
1000100011100000?120010010211421001?100-
10 ? 100000001701010000000000 ? 0 ??? 00001 ???-
000000? 12?000011110?001000?0?000?00??0??0?
Kayenta Form
2 ? 1 ???? 1 ? 0 ??? 0 ? 1 ?? 1 ???? 11 ?????????? 2171 ( 01 ) 10 -
0020 ? 1 ? 011 ? 1100 ? 10 ???????? 10171 ? 000 ?? 10 ????-
???0?????????????? 1 ??????? 1 ?????00100??????00??-
10?00??011211 ??001 ?????0?0?00???? 1011111??-
0? 110??0??? 1 ?????????? 0?
Zosuchus
201 ?? 0 ? 1200000 ?? 001010 ( 01 } 1107001110 ? 022 -
1101001271 ?? 01171100070 ? 1 ? 0211110 ???? 0?01 -
111 ???????????????????? 1 ? 12 ? 3 ???? 1 ????? 001000 -
11011 ?0001 ?0?0010112?(01 )?0001???0?00???01 -
0111 ?? 1011710111000000100 ??? 0 ? 000?00
Fruita Form
201 ?? 001200100010000100100000110010221 ?-
11 ? 0020112 ? 1 ??? 0 ? 0 ?? 0 ?? 1 ? 2 ? 31 ????? 1 ? 011110 -
1011 ? 1 ? 00011112 ? 0 ?? 1 ??? { 01 } 00 ??? 1 ? 1001 ? 00 -
1 ? 0 ? 0100100 ?? 101 ? 0011701110 ?? 0 ?? 00 ? 107000 -
0 ? 1 ??? 000 ???? 101 ? 0 ? 00000 ? 000 ?? 0 ??? 0??0
Hsisosuchus
(12)01110?1200000? 10010?0??00????0?0??? Ill- 211??????1??000000100001100011000?022110-
107002010011 111 10000101172011?0102100?1- 1000(12)?? 12? 11 ? 1000070? 1 ?0?? Ill ?4?00(01)0-
38
AMERICAN MUSEUM NOVITATES
NO. 3512
2717710777777700071000777101700217717777701-
GO 17777777000077007? 1711717700777777770770777-
{01} 007070111(01 )?00??00?0? 10007070??? 10
Notosuchus
10170011010100111000111111001100010221101-
10021112011710000107110211111270101110001-
{01} 1117172000010007701220117771100101(01)1-
101(01)0100100000011111111117000111100100-
00101110110000111011000000000000000001111
Comahuesuchus
103770? 10177007777701127777777700 1 0727??? 1 ??-
1171 ????????????????? 1317777770? 10101 ?????????-
?????????????? {01} 1377? 17777770? 1071012017017-
7777011770? 1 ???? 1177711700 1 0071707700077? 10-
07707000770777701111071
Mariliasuchus
101700710100001110001(01)0111000110010221-
10100021712011710000107170213111210(01)07-
11000?????????????????????2211177711?????0{01 }0-
00101001071100{01 }00111101107001177701100-
0100110010000171010000000000077701111071
Uruguaysuchus
201700110177007? 107? 1771 ????177701022? 1017-
0011 ???? 177777077077701111(12)777000110 1 007-
717177777700007077017210021007007000? {01} 7-
7701717007??? 170111711 ????? 11 ????? 170001 ????-
7777077? 1077777007777777707777? 17
Chimaeresuchus
10170001111700???????????????????????????????? -
??????????????????? 127701107010107 ? 171 ?????21 -
00700???? 11 (12)731421077007010011111101177-
7777070110?????????? 10711 ?????3 ??????????? 1700-
777077777777770707777?
Malawisuchus
10170071110000?(01)10001(01){01} 110071100-
017221101000117720???1000? 1071702? 111(01)-
270101110001 ???? 1772100000107701(12)211177-
70170777011001017110007771101101017070001-
7770? 100772111071000011100000000000007770-
7700710
Candidodon
??????????????????????????????????????????????????-
??????????????????????????????????????????????????.
??????????????????? 1 ????????????????0??0????????7-
?????? 1 ????2???????????????????????????????????7?
Simosuchus
103010110000001000101111107011000102171-
010001171101171000010717020112121010110-
00077777770210072010? 1000201077701 ?????? 11 -
011012120000101001110021100120777211{12-
}0001111011001{01}171000000000001000007-
000710
Sphagesaurus
101700010177007? 100????? 1107777777772110170-
07777701171000???????? 1372???????? 100????????-
I ???????????????3 1277770??????? 1111110111 lll-
II 1111111001110111 170711707701170?1077017-
7000000700777777700707? 17
Bretesuchus
1 {01} 077011217700???????????0??????????2?7? 10-
011 ???????? 101171 ?????? 13? 17? 1700? 10110?????-
?????????????????? 100777? 1777777701770777701770-
77707? 170777???????? {01} 0(01)? 1710? 177? 177001 -
??00??????????0????0?01717
Baurusuchus
100770712177007110177771117011077777271011-
0011112011710007107710773111210101111117-
???????????????????? 12103777? 1 ????? 1101110101-
01110011001111011070111???{01}0(01)11110-
1110170000170001000000000077707701111
Iberosuchus
1707000127070011100011111170170007027?10-
1001117127? 171017? 10? 1 ???? 1117? 1070? 101101-
1 ?????? {12} {1234} 00770077700? {12} (01 )0?2??0-
000777110011010107170771007110017077101777-
{01 }?011170011017077017100001000000770770-
700771?
Araripesuchus gomesii
201000110100001110001011111011(01)00102-
211010001111201171000070711020112121000-
1101{01}{01}1{01}11111?1{234}0001000100-
11110021001001010100100100100000010011-
00021000011070011{01}000011110100001110-
07000000000000007000710
Araripesuchus patagonicus
2010007101000071{01}0001011117011100102-
27101000117127117100077071702711212707011-
2006
ZAHER ET AL.: CRANIAL MORPHOLOGY OF MARILIASUCHUS AMARALI
39
{01}1??1?1 ??????????1000??01111007770177707-
0170110170100007? 100110102707701 ????0?? {0 -
1}1000111701000011107000000000000000700-
0710
Lomasuchus
Pelagosaurus
20271111110011020101000000000000(01)100-
211010000001101111001001710001200730000-
0200001101117000000120001110170077? 10777-
???? 171 ??????0000??01001070010???00????0001 -
0000112010000110000017000000077707700? 10
201 ???? 121170071100010111177110001022710-
100017? 127? 171007?17717727217777700770(12)1-
1 ???????????????????? 1777007770077777070077? 171-
1070077700011707? 17707777770107770? 117710770-
171000771100077077707070? 10
Peirosaurus
20170117? 17700?????? 10? 17777777707772? 10?????-
?????????????????????? 1 ??????????(12)1 ???????????-
?????????????000????????????0??7? 1777077777770? 1 -
??????????????(01 )??????0??????????????007? 1 ????-
7777770???????
Theriosuchus
20110111110100110000110111100110011721-
1010001711701111000????? 1720211741001010-
10110111110001111200100101000270071071-
10110(01)0017110070070700 1 0077017707007? 1 -
010000071170107701710000??0000??????0???0-
710
Alligatorium
Teleosauridae
(02)02?11111100110201001000000000001100-
2170100070011011110010117170012000370007-
20000210111170000? 120001010117077? 107701 -
0011017? 1011000011000010100707700007? 100-
0100001170107011100010100000000000077007-
10
Metriorhynchidae
(02)02712110100111201011000700000001100-
2170700070011011110010117170012007300010-
200002101711700007777770770127077? 10070100-
11017? 10117000770000101027077700077770010-
000117010007110000010000000077700700? 10
Sokotosuchus
27277? 11017710777700 1 00177? 1010017012? 1 ????-
7? 111271171711770??? 1717077777777701 ?????????-
??????????????????????????????? 177777777777777770-
??0????????????????????0????????????????????????7-
????????????
?0?????? 170000? 10000107111770? 100? 177770770-
07? 117? 17? 1000777777720? 1777700 1 01710170111-
1100077? 1700100??????????? 107? 1 ???????????????-
????0???????????????????????0??????????????????07-
?????????????????
Goniopholis
203712111100101110001001117001000100271-
01000?Ill201171010?10?1 ?02131274100{01)0-
(12)0201171771770700?1200?117000002100010-
7110110177101100700001001000171777000011-
002000001100100001111070100000000000070-
00710
Dyrosauridae
0027? 171017010? 1170010001171010011012? 10-
1010011120117101171071010130273700772700-
0777777770700?????? 1 ??????????????????? 1 ????????-
????????0??0???????????????02170001777707777770-
07? 17000000077707700? 10
Pholidosaurus
212? 111101771171110110011170010001072117-
10000111211171017? 10? 100? 1311730077727077-
71171777077077270????????????????????? 1 ???? 1711-
0777770700 1 07777777777777770? 17000177? 1070017-
10070 1 0777077777707770? 10
Eutretauranosuchus
Bernissartia
2037777171001011100010011170070001001110-
7000? 111201171010770? 170?121204700001020-
11177? 1770770? 1 ?????????000???00?????0? 101 ???-
711077777770770077? 17770777? 10727770017077000-
7171107017000000077707007? 10
2037721111770011100070011170010007002????-
7000111271171010070? 177? 17177410010102011-
71711770200211101101000007700???????? 1 ????-
1777777770770? 10777017770777? 1712000001777707-
77777007? 100 00777707707770? 10
40
AMERICAN MUSEUM NOVITATES
NO. 3512
Hylaeochampsa
00??????? 11 ??? 11 ???? 1 ?01 ???0????0?002? 1 ? 1011 -
????????? 101 ?? 1 ?? 1 ????? 10???????????????????????-
???????????????????0??????? 10????????????0??0???-
???0????????????2??00????????????????????????????-
?0???0?1?
Borealosuchus
203?1211110010111000100111?001000100211-
010111111211111010010?110?1310031000110-
?011111111113111?110?00?000002110?100100-
?101??11110??000000010001?1???0000110?20?-
00011001000011100001000000000000 ??? 0 ? 10
Gavialis
212 ? 12111100111111011011111001000100211 -
0101101112011110110101110{01}131003100-
0120000011 111 10131112111100?000002110?1 -
00100 ? 101 ?? 121100 ? 00000001000101 ? 1 ?00001 -
? 0 ? 20 ? 00011001000011100001 ? 00000000000 ??-
00?10
Crocodylus
203012111100(01)0111000102111100100010021-
10?01111112011110100101110{01}13100310001-
0010121111110131112021100?00000211001001-
00 ? 101 ?? 121100 ? 0000000100110101100001 ? 0 ? 20 -
000011001000011100001000000000000 ? 000?10
Alligator
203112?101?0001110001021111001000?0021101-
01111112011110100101110{01}03120310001002-
01211111111311120211?0?000002110010010011-
01??111000?00000001001(12)01011000011(01)12-
00000110010000111000010000000000000000?10
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